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LBERT R. MANIV.
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CORNELL UNIVERSITY LIBRARY
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COTAXON
AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION
OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS
Voiune XXVII October-December 1986 ae
CONTENTS
Studies on ectomycorrhizae III. Mycorrhizae formed by four fungi
in the genera Lactarius and Russula on spruce .............. R AGERER:
A new species of the genus Oidiwmn from India.
Coniochaeridium mirabile, a new ascomycete isolated from salted food.
SHUN-ICHI UDAGAWA AND HARUO TSUBOUCHT a
Materials for a lichen flora of the Andaman Inslands — I.
M. B. NAGARKAR, P. K. SETHY, AND P. G. PATWARDHAN a ee,
Hyphomucor, anew genus in the Mucorales for Mucor assamensis.
Metarhizivn flavoviride var. minus, var. NOv., a ‘a pathogen of plant- and
leafhoppers on rice in the Philippines and SolomonIslands.
MICHIEL C. ROMBACH, RICHARD A. HUMBER, AND DONALD W. ROBERTS 87 ee
Studies on Chinese ascomycetes. 2 Astrosphaeriella lageniformis.
JING-ZHU YUE AND OVE ERIKSSON: 93 x. ae
3 hy oieipara anew thermotolerant genus of Peo ceonS
hyphomycetes.
SHUN-ICHI UDAGAWA, TAKEYOSHI AWAO, AND SAMIR K. ABDULLAE 0 |
Gibbago, a new phaeodictyoconidial genus of hyphomycetes.
A new species of the lichen genus Sulcaria (Ascomycctina, Alectoriaceae) from
BE TER C ors hls BUS RABIN eh etd ou Fale Moni oath MOEN a ale Gaus . IRWIN aS BRODO ie e : ae
Badarisama soja, a new bulbil-forming genus and species.
I. K. KUNWAR, J. B. MANANDHAR, AND J. B. SINCLAIR. Cas
Notes de nomenclature concernant les Hyménomycétes; IV. Sur quelques
épithetes spécifiques preoccupés. 3°... ce eek: REGIS COURTECUISSE 127 en
Two new dematiaceous hyphomycetes from tropical forest litter.
LAURA ZUCCONI AND SILVANO ONOFRI 147°
| Digitatispora lignicola, a new marine lignicolous Basidiomycotina.
Marine fungi from Seychelles. VI. Massarina velataspora, a new marine
G. BAGYANARAYANA AND U. BRAUN, oe a
M.A. A. SCHIPPER 83. a ae
i .
|
EMORY G. SIMMONS 107. ee
E. B. GARETH JONES 155. . oe
Ascomycotina from mangrove wood........ K. D. HYDE AND B, D. BORSE 161
Taxonomic concepts in the Endogonaceae: III. The separation of Scutellospora
gen. nov. from Gigaspora Gerd. & Trappe.
CHRISTOPHER WALKER AND F. E. SANDERS: 169
[CONTENTS continues overleaf]
ISSN 0093-4666 MYXNAE 27 1-628 (1986)
Published quarterly by MYCOTAXON, Ltd., P.O. Box 264, Ithaca NY 14851
For subscription details, availability in microfilm and microfiche,
and availability of articles as tear sheets, see back cover.
[CONTENTS continued from front cover]
Five new species of Parmelia (Lichenes, Parmeliaceae) from southern Africa.
FRANKLIN A. BRUSSE
Hypogeous fungi from the southeastern United States I. The genus Rhizopogon.
STEVEN L. MILLER
Species of Scutellospora (Endogonaceae) with smooth-walled spores from
maritime sand dunes: two new species and a redescription of the spores of
Scutellospora pellucida and Scutellospora calospora.
R. E. KOSKE AND CHRISTOPHER WALKER
Four new effigurate-crustose species of Parmelia (Lichenes, Parmeliaceae) from
SOWLMO MI AIICA 6h. Sec os tinie WARN & ini tee aeeee as FRANKLIN A. BRUSSE
Bertiella (Sacc.) Sacc. & Sydow a synonym of Massarina Sacc.
OVE ERIKSSON AND JING-ZHU YUE
A new Chlamydomyces species from Italy ................ CLAUDIA PERINI
Notes on Entomophthorales (Zygomycotina) collected by T. Petch: I. Erynia
ellisiana sp. nov., non Erynia forficulae (Giard), comb. nov., pathogens
ol Poriculidae (Dermaptera) 00's 05. wikis tie sae ts ISRAEL S. BEN-ZE'EV
Canoparmelia, Paraparmelia and Relicinopsis. Three new genera in the
Parmeliaceae (lichenized Ascomycotina).
JOHN A. ELIX, JEN JOHNSTON, AND DOUGLAS VERDON
EWG) NEW GISCOIY COLES ONG INES, O65. is vars wistetectiesuece in ys ees A. FUNK
A comparison between Phytophthora cryptogea and P. drechsleri.
H. H. HO AND S. C. JONG
Tremellostereum (Tremellaceae) nov. gen................-4- L. RYVARDEN
Basidiomycetes that decay Gambel oak in southwestern Colorado: III.
J. PAGE LINDSEY
Lahmia Korber (= Parkerella A. Funk) a misinterpreted genus with isolated
PRIS SENOS hie et cng 9 Ria et Lalas 12, rag aes nena ele eA OVE ERIKSSON
Cultural studies in Tubulicrinis and Xenasmatella (Corticiaceae, Basidiomycetes).
NILS HALLENBERG
Foliicolous ascomycetes 7. Phylogenetic systematics of the Capnodiaceae.
DON R. REYNOLDS
NOTICE: Guide: to Sanctioned Fungal Naines . 26.005)5 sav ais Sta oe ie ee
The use of isozyme analysis in fungal taxonomy and genetics.
J. A. MICALES, M. R. BONDE, AND G. L. PETERSON
The lichen flora of the Galapagos Islands, Ecuador........ WILLIAM A. WEBER
Platygloea acanthophysa, a new species with single sterigmate basidia and
ACANUMODNVSES wis %, Ae tees pie sos oe BS: hoe HAROLD H. BURDSALL, JR.
Macrochemical color reactions of macromycetes II. Chemical properties and
systematic position of Bondarzewia mesenterica ....... HARALD M. FRANK
East African rusts (Uredinales), mainly from Uganda 5. On families belonging
to'Gamonetalae:.) edaiate ss axcinlces ge oles re oe te 4 HALVOR B. GJERUM
Confirmation of Ascosphaera apis in Georgia.
R. T. HANLIN AND R. A. SAUNDERS
Récoltes de Biscogniauxia dennisii (Pyrenomycetes, Ascomycetes) dans le
Sud -onestice la Francés 26 os ee cles 3 ee eee FRANCOISE CANDOUSSAU
New species of the lichen genus Xanthoparmelia from southern Africa
(Ascomycota: Parmeliaceae) ..6' 65 06. 5.0,+ 3 saree i MASON E. HALE, JR.
New species of Relicinia (lichenized Ascomycotina) from Australasia.
JOHN A. ELIX AND JEN JOHNSTON
[CONTENTS continues inside back cover]
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255
263
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MYCOTAXON
AN INTERNATIONAL JOURNAL DESIGNED TO EXPEDITE PUBLICATION
OF RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI & LICHENS
ALBERT R. MANN
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DEC 1 ‘7 1986
ITHACA, N.Y. 14852
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CO-EDITORS
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Laboratoire de Mycologie systématique et appliquée
Université de Louvain, B-1348 Louvain-la-Neuve, Belgium
RICHARD P. KORF
English Language Editor & Managing Editor
Plant Pathology Herbarium, Cornell University
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TABLE OF CONTENTS, VOLUME TWENTY-SEVEN
October-December 1986
Studies on ectomycorrhizae III. Mycorrhizae formed by four fungi
in the genera Lactarius and Russula on spruce .............. R. AGERER 1
A new species of the genus Oidiwmn from India.
G. BAGYANARAYANA AND U. BRAUN 61
Coniochaetidium mirabile, a new ascomycete isolated from salted food.
SHUN-ICHI UDAGAWA AND HARUO TSUBOUCHI 63
Materials for a lichen flora of the Andaman Inslands — I.
M. B. NAGARKAR, P. K. SETHY, AND P. G. PATWARDHAN 71
Hyphomucor, a new genus in the Mucorales for Mucor assamensis.
M. A. A. SCHIPPER 83
Metarhizium flavoviride var. minus, var. nov., a pathogen of plant- and
leafhoppers on rice in the Philippines and Solomon Islands.
MICHIEL C. ROMBACH, RICHARD A. HUMBER, AND DONALD W. ROBERTS 87
Studies on Chinese ascomycetes. 3. Astrosphaeriella lageniformis.
JING-ZHU YUE AND OVE ERIKSSON 93
Thermophymatospora, a new thermotolerant genus of basidiomycetous
hyphomycetes.
SHUN-ICHI UDAGAWA, TAKEYOSHI AWAO, AND SAMIR K. ABDULLAH 99
Gibbago, a new phaeodictyoconidial genus of hyphomycetes.
EMORY G. SIMMONS 107
A new species of the lichen genus Sulcaria (Ascomycotina, Alectoriaceae) from
MPLOTIsaY kor eect Pua eH Ee be eitcces CHAD eat IRWIN M. BRODO 113
Badarisama sojae, a new bulbil-forming genus and species.
I. K. KUNWAR, J. B. MANANDHAR, AND J. B. SINCLAIR 119
Notes de nomenclature concernant les Hyménomycétes; IV. Sur quelques
épithétes spécifiques préoccupés.3............... REGIS COURTECUISSE 127
Two new dematiaceous hyphomycetes from tropical forest litter.
LAURA ZUCCONI AND SILVANO ONOFRI 147
Digitatispora lignicola, a new marine lignicolous Basidiomycotina.
E. B. GARETH JONES 155
Marine fungi from Seychelles. VI. Massarina velataspora, a new marine
Ascomycotina from mangrove wood........ K. D. HYDE AND B. D. BORSE 161
Taxonomic concepts in the Endogonaceae: III. The separation of Scutellospora
gen. nov. from Gigaspora Gerd. & Trappe.
CHRISTOPHER WALKER AND F. E. SANDERS 169
Five new species of Parmelia (Lichenes, Parmeliaceae) from southern Africa.
FRANKLIN A. BRUSSE 183
Hypogeous fungi from the southeastern United States I. The genus Rhizopogon.
STEVEN L. MILLER 193
Species of Scutellospora (Endogonaceae) with smooth-walled spores from
maritime sand dunes: two new species and a redescription of the spores of
Scutellospora pellucida and Scutellospora calospora.
R. E. KOSKE AND CHRISTOPHER WALKER 219
Four new effigurate-crustose species of Parmelia (Lichenes, Parmeliaceae) from
MOMIEMC TIL GALLIC A 1th. co) St.0 die yeils sce ies Marerebabe aieew cated FRANKLIN A. BRUSSE 237
Bertiella (Sacc.) Sacc. & Sydow a synonym of Massarina Sacc.
OVE ERIKSSON AND JING-ZHU YUE 247
A new Chlamydomyces species from Italy ................ CLAUDIA PERINI 255
Notes on Entomophthorales (Zygomycotina) collected by T. Petch: II. Erynia
ellisiana sp. nov., non Erynia forficulae (Giard), comb. nov., pathogens
Gmrormicnlidae (Dermaptera). i.e) ccs Gee Wt ckdls tors ISRAEL S. BEN-ZE'EV 263
Lv
Canoparmelia, Paraparmelia and Relicinopsis. Three new genera in the
Parmeliaceae (lichenized Ascomycotina).
JOHN A. ELIX, JEN JOHNSTON, AND DOUGLAS VERDON
Two mew. discOmycetes On Pinus. Oo a0). cc oe a ce ee A. FUNK
A comparison between Phytophthora cryptogea and P. drechsleri.
H. H. HO AND S. C. JONG
Tremellostereum (Tremellaceae) nov. gen...............202. L. RYVARDEN
Basidiomycetes that decay Gambel oak in southwestern Colorado: III.
J. PAGE LINDSEY
Lahmia Korber (= Parkerella A. Funk) a misinterpreted genus with isolated
POSIELOMES F cence Punts Oh Une meeeent citiadare | ae enn ke Ran nee OVE ERIKSSON
Cultural studies in Tubulicrinis and Xenasmatella (Corticiaceae, Basidiomycetes).
NILS HALLENBERG
Foliicolous ascomycetes 7. Phylogenetic systematics of the Capnodiaceae.
DON R. REYNOLDS
NOTICE: Guide to’Sanctioned Fungal Names'.."'20\". so... ene eee ee
The use of isozyme analysis in fungal taxonomy and genetics.
J. A. MICALES, M. R. BONDE, AND G. L. PETERSON
The lichen flora of the Galapagos Islands, Ecuador........ WILLIAM A. WEBER
Platygloea acanthophysa, a new species with single sterigmate basidia and
acanthophyses noc. Acasa ie Wee ee al HAROLD H. BURDSALL, JR.
Macrochemical color reactions of macromycetes II. Chemical properties and
systematic position of Bondarzewia mesenterica ....... HARALD M. FRANK
East African rusts (Uredinales), mainly from Uganda 5. On families belonging
COLGaMOpetalac irs. lee ices tice bes ares Peace HALVOR B. GJERUM
Confirmation of Ascosphaera apis in Georgia.
R. T. HANLIN AND R. A. SAUNDERS
Récoltes de Biscogniauxia dennisii (Pyrenomycetes, Ascomycetes) dans le
slid-ouest. dé lalFrance. 7.50.2 a ee ee FRANCOISE CANDOUSSAU
New species of the lichen genus Xanthoparmelia from southern Africa
(Ascomycotina:-Parmeliaceae) .. 2000s noe es ee MASON E. HALE, JR.
New species of Relicinia (lichenized Ascomycotina) from Australasia.
JOHN A. ELIX AND JEN JOHNSTON
‘AUINOLSINDE Kies he, Ne tee Pes alan hw bie a cre creo enon Guero tuiaae oe tae a ee
INDEX to Fungous*and Lichen Taxa"... / 200° i, 3.) ee ne eee ee ee
RREVICWETS & FHA 8 Oe, cree os a ee Ie ae on ene ee
Publication date; MYCOTAXON Volume 26! 0703. 2 vs cee oa ee eee
Errata: eiessd eS ae eB a Ee PIS a tae) ey ene
241
283
289
321
325
347
361
377
404
405
451
499
503
507
od
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MYCOTAXON
McnieerXNY LL yt Diet oo October-December 1986
Studies on Ectomycorrhizae III *
Mycorrhizae Formed by four Fungi in the
Genera Lactarius and Russula on Spruce
R. AGERER
Institute for Systematic Botany, University of Munich
D-8000 Munich
Menzingerstrafke 67
This series describes mainly anatomical and morphological
studies on ectomycorrhizae. Keys to the characterized
mycorrhizae will be provided. In later papers of this
series, the keys will be enlaraed by inclusion of newly
described mycorrhizae. In part II of the series the
methods of characterization and identification applied in
this series are described comprehensively.
SUMMARY
The anatomical and morphological characteristics of four
identified ectomycorrhizae are described. The mycorrhizae
of Lactarius deterrimus, L. picinus, Russula_ ochroleuca
and R. xerampelina were identified by tracing hyphal
connections between the fruitbodies and the mycorrhizae.
The four mycorrhizae are characterized by strikind
differences. Besides variations in the structure of the
mantle, there are differences of colour and shape of the
mycorrhizae, and of their attached rhizamorphs, different
colour reactions with reagents, and differences regarding
laticiferous hyphae and Hartiq net. All mycorrhizae belong
to the so-called smooth type. Their features are compared
with those described in the literature as being typical of
these or related species.
* AGERER, R. (1986) - Studies on Ectomycorrhizae
II. - Introducing remarks on characterization and
macntification. Mycotaxon .26:) 473, ~,4927
KEY WORDS
ectomycorrhizae-characterization, -identification, -anatomy,
-morphology, -surface view, -longitudinal section, -cross
section, inner surface of mantle, chemical reaction,
autofluorescence, nucleus, siderophilous granulation,
emanating hyphae, rhizomorph, cystidia, laticiferous hyphae,
Picea abies, Lactarius deterrimus, L. picinus, Russula
ochroleuca, R. xerampelina, key.
I KEY FOR THE CHARACTERIZED MYCORRHIZAE
1 Mantle of mycorrhizae with laticiferous hyphae
2 Surface of mantle with normal hyphae running
more or less parallel to the surface of the
mycorrhizae
Lactarius deterrimus
2 Surface of mantle with short, obtuse hyphal
ends protruding more or less perpendicularly
to the surface of mantle
Lactarius picinus
1 Mantle of mycorrhizae without laticiferous
hyphae
3 Surface of mantle pseudoparenchymatous, cells
anqular
Russula ochroleuca
3 Surface of mantle a network of hyphae
Russula xerampelina
IT DESCRIPTIONS OF THE MYCORRHIZAE
1. LACTARIUS DETERRIMUS Groger
FAG lipuie poe oipauee we lal gine
MORPHOLOGICAL CHARACTERS
Ramification: monopodial; length of unramified
ends: up to 2,1 mm; diametre of unramified ends:
0,32 - 0,52 (0,65) mm; shape of unramified ends:
straight or slightly twisted, sometimes more or
less flat; structure of surface: even, with some
rhizomorphs.
Colour of unramified ends: bright copper-orange;
colour of tips: bright copper-orange; colour of
older parts: brown with dark green patches.
ANATOMICAL CHARACTERS OF SURFACE
OUTER SURFACE OF MANTLE
Structure: plectenchymatous, individual hyphae
recognizable; size of hyphal cells: (2) 3 - 4 pm;
thickness of cell walls: not conspicuously
thick-walled; surface of the very tip:
plectenchymatous, hyphae strongly agqqlutinated, 2
- 3 (3,5) pm thick; laticiferous hyphae, 3 - 7 pm
in diam., with 0,5 - 1 pm thick walls, contents
yellow.
RHI ZOMORPHS
Shape in cross-section: more or less round;
arrangement and diameter of hyphae: hyphae more or
less parallel, without thicker central hyphae, 2,5
oye An, Olam. , rarely with, laticiferous
hyphae, scarcely thicker than the other hyphae;
thickness of cell walls: not conspicuously
thick-walled; colour of hyphae: pigment
membranaceous, slightly yellowish; septae: without
clamp connections; anastomoses: rare, hyphae
aqglutinated.
EMANATING HYPHAE
Shape and diameter: more or less straight, ca. 2,5
pm in diam, seldom to be found; thickness of cell
walls: ca. 0,5 pm; colour of hyphae: more or less
colourless; septae; without clamps.
CYSTIDIA none found.
INNER SURFACE OF MANTLE
Arrangement of hyphae; irregular, hyphae short or
longer; diameter of hyphae: (2) 3 - 4 (7) pm.
ANATOMICAL CHARACTERS, CROSS-SECTION
MANTLE
Thickness and differentiation: (10) 20 - 30 (40)
pm, strongly agglutinated, hyphal walls strongly
swollen; cell-shape and dimensions of different
layers: no differentiated layers recognizable,
hyphae tangentially 2 - 10 (15) pm, radially 1 -
2,5 pm, laticiferous hyphae tangentially 4 -
20(40) pm, radially 4 - 5 pm.
TANNIN CELLS
Number of rows: 1 - 2; shape and measure: narrow,
tangentially oriented, 30 - 40 x 5 - 10 pm.
HARTIG NET
Shape and dimensions of cortex cells:
tangentially—oval,! 30°'- 60° x (10)° 15 =" 20 (30) am;
depth of H'net: 3 rows including tannin cells, one
row next the endodermis being free of H'net;
thickness of H'net: (5) 6 - 7 (8) pm; number of
hyphal rows of H'net: 1 ; shape of H'net cells in
section: oval, but often distinctly stretched.
ANATOMICAL CHARACTERS, LONGITUDINAL SECTION
MANTLE
Shape and dimensions of the cells of the very tip:
like the hyphae of the remaining part of the
mycorrhiza, but hyphae less agglutinated; shape
and dimensions of cells in different layers: no
distinct layers recognizable, hyphae tangentially
2. —. 10" (20) pm, radially 2.-)3 (5) °pm.
HARTIG NET
Shape and dimensions of cortex cells: flat rhombic
to oval, more or less parallel to the axis of the
mvcorrniza,, 000 — 110 x12 —) 20 (30) pam ; shape and
dimensions of tannin cells: flat, parallel to the
axisvot the mycorrhiza, 50-90 (120). 3 ae
inner tannin cell layer up to 15 pm thick.
COLOUR REACTION IN DIFFERENT REAGENTS
Acetic-Fuchsin: ----; a-Naphtol: ----};
Ammonium-Hydroxide: ochre-orange, especially
laticiferous hyphae; Brilliant-Kresyl-Blue:
bluish-green, laticiferous hyphae conspicuous,
after treatment with water violet blue,
laticiferous hyphae distinctly green;
Chlorazol-Black E: slightly bluish- green;
Cotton-Blue: slightly blue; Erythrosin: reddish,
especially thin laticiferous hyphae; Fast-Green:
green, especially laticiferous hyphae; Gujak:
Fig. 1 - Lactarius deterrimus - a: Habit of
mycorrhizae. - b, b', b'': different surface
views. - c, c',c'', c'''s: inner surface of mantle,
different views. - d: Surface view of the very
tip. (AL Letigs. from 072028) sin Herb. R. Agerer..
--—-3 Iron-Sulfate: ----; KOH 15%: ochre-orange,
especially laticiferous hyphae; Kongo-Red:
slightly blood-red; Magnesium-Acetate: ----;
Malachit-Green: green, especially laticiferous
hyphae; Martius-Yellow: ----; Melzer's Reagent:
----; Phenol: ----; Phenol-Anilin: ----; Phloxin
B: reddish, especially laticiferous hyphae;
Stahl's EP-Reagent: ?; Sulfo-Vanillin:
laticiiferous hyphae with dark granules;
Toluidin-Blue: violet blue.
AUTOFLUORESCENCE
WHOLE MYKORRHIZA
UV 254 nm: ----; UV 366 nm: ----.
MANTLE AND RHIZOMORPHS
UV-filter 340 - 380 nm: blue, laticiferous hyphae
distinctly blue; BLUE-filter 450 - 490 nm:
greenish, laticiferous hyphae distinctly green.
STAINING OF NUCLEI
Hyphal cells of outer mantle surface with two
nuclei lying mostly close together, seldom hyphal
cells with 3 or 4 nuclei (method: aceto-carmine).
Siderophilous reaction negative.
MATERIAL STUDIED AND METHOD OF IDENTIFICATION
Germany, Bavaria, Miinchen-Oberschleif8heim,
Korbiniani-H61zl, Versuchsflache O, Quadrat 2028,
1267. 1983s Herb. RA 2028 C247). 83.) wichericaa
abies (material for microscopical studies). -
Germany, Bavaria, Munchen-OberschleifBheinm,
Korbiniani-H6l1zl, bei Versuchsflache O, Herb. RA
Oa(22.9.84), with Picea abies (material for
staining). There were direct connections between
Fiq. 2 - Lactarius deterrimus - a: Surface view of
laticiferous hyphae. - b, b', b'': Rhizomorphs
(arrows indicate the distal direction). - c:
Rhizomorph of basis of fruitbody. (Fidqs. a - b''
from 0/2028 in Herb.’ R.), Agerer,, fig. -c) from, Ca
(22. 9. 84) in Herb. R. Agerer).
mycorrhizae and the rhizomorphs of the base of
fruitbody.
COMPARISON OF THE MYCORRHIZA OF LACTARIUS
DETERRIMUS WITH THOSE OF L. PICINUS
(Selected features only)
The mycorrhizae of Lactarius deterrimus differ
considerably from those of L. picinus.
(1) Morphological characters: Whereas L. picinus
forms pyramidal, densely ramified mycorrhizae,
those of L. deterrimus are mostly more or less
pinnate and the individual mycorrhizae are
relatively scattered. Additionally the terminal
portions of the mycorrhizae are much longer in L.
deterrimus than in L. picinus,. Besides the
different colour (brightly copper-orange with
olive-green patches in L. deterrimus, fleshy-brown
in L. picinus) the gross morphology of the
surfaces deviates fundamentally. The surface of
the mycorrhiza of L. picinus is pruinose, that of
Le. Geterrimus is smooth.
The most obvious structural differences
between the two mycorrhizae are found in the
mantle. In L. deterrimus the mantle surface is
composed of more or less undifferentiated hyphae
whereas those of L. picinus form special endcells
which project more or less perpendicularly to the
surface. A second difference regards the diameter
of the laticiferous hyphae: in L. picinus they
are distinctly broader than in L.deterrimus.
Moreover, sectioned mycorrhizae of L. picinus show
a three-layered mantle; that of L. deterrimus is
more or less homogenous. The rhizomorphs of L.
picinus possess broader hyphae in the centre than
in the periphery, those of L. deterrimus are of
Fig. 3 - Lactarius deterrimus - a: Habit of
mycorrhiza and fruitbody primordium. - b: Surface
view. - c: Surface view of the very tip. - d:
Surface view of laticiferous hyphae. - e, e':
Inner surface of mantle, different views. - f:
Rhizomorph of the basis of fruitbody. - gq:
Emanating hyphae. (All figs. from Oa(22. 9. 84) in
Herb. R. Agerer).
10
of more or less uniform hyphae, with septae at
short intervals.
Within the cortex, cells of the Hartig net in
L. picinus are thinner than in L. deterrimus.
(2) Characters after treatment with different
reagents: In alkalies (KOH and Ammonium-hydroxide)
the mantle hyphae of Lactarius deterrimus become
orange-ochre, especially the lactiferous hyphae
whereas L. picinus do not show such reactions.
Another difference is due to Iron-sulfate: only
the mycorrhiza of L. picinus turns greenish-brown.
Third, only in L. picinus does Melzer's reagent
tinge the mantle yellowish-brown, especially the
laticiferous hyphae. There are some other slight
differences in color-reactions, but they are not
disctinctive enough to differentiate these two
species. We found no mention of colour reactions
of L. deterrimus with the above mentioned reagents
in literature (SCHMITT 1973, MOSER 1978). Only
ROMAGNESI (1958) mentions for L. deterrimus (=
Lactarius deliciosus var. picinus) a reaction with
Gujak. Unfortunately we did not check this reagent
with fresh mycorrhizae of L. deterrimus.
MYCORRHIZAE OF LACTARIUS DETERRIMUS AS TREATED IN
LITERATURE
This species was described rather recently (GROGER
1968), and no information about its mycorrhizae is
available. Of the section Dapetes only L.
deliciosus and L. sanguifluus is dealt with in the
literature. But the researches on this species
probably refer to more than L. deliciosus s. str.
Pl. 1 - Lactarius deterrimus -a: Habit of
mycorrhizae. - b: Autofluorescence of laticiferous
hyphae (Blue-filter). - c: Inner surface of
mantle. - d, e: Rhizomorphs. (Figs. a, d, e from
0/2028 in Herb. R. Agerer, figs. b, c from Oa(22.
9. 84) in Herb. R. Agerer). - Bar 1 mm (fig. a),
LOM figs bo e).
\ (
at
\
LW
12
Similar, closely related species in this group
were possibly included, because the species
concepts had not been well defined untill
recently.
HATCH & HATCH (1933) isolated cultures of
Lactarius deliciosus from fruitbodies and
synthesized mycorrhizae with Pinus species. They
noted that the most abundant associated trees were
Pinus and Betula species; Lactarius deliciosus is
known as a species associated with pines but not
of Picea , in contrast to L. deterrimus. The
authors confirmed the ability of L. deliciosus to
form mycorrhizae but provided no detailed
description showing only a longitudinal section
through the mycorrhiza. A plectenchymatous mantle
and a Hartig net including up to three rows of
cortex cells were characterizing features.
RIFFLE (1973) isolated cultures from sporocarps of
Lactarius deliciosus growing with Pinus ponderosa
and synthesized mycorrhizae with seedlings of this
tree. Without doubt, this fungus was correctly
identified, because Pinus was the host. The
pictures of the sectioned mycorrhizae resemble the
above described mycorrhiza of L. deterrimus. The
mantle is plectenchymatous. RIFFLE (1973)
mentions, however, that in his cultures no
rhizomorphs grew. This may be due to the high
nutrient status of the agar medium used for
synthesizing mycorrhizae. The colour of the
synthesized mycorrhiza of L. deliciosus was cream
to tan and not brightly copper-orange. The
dichotomously forked mycorrhizae of L. deliciosus
+ Pinus ponderosa are typical of pine species
(BRUCHMANN 1874, MELIN 1923, MODESS 1941, SLANKIS
1948).
Pl. 2 - Lactarius deterrimus - a, b: Cross
sections. - c,d: Longitudinal sections. (All figs.
from 0/2028 in Herb. R. Agerer). - Bar for all
figs. 10 pm.
13
14
ASCAI & LARGENT (1983) found fruitbodies of
Lactarius deliciosus s.l. associated with Abies
concolor but it is questionable if this is the
species in the restricted sense. In Europe
Lactarius salmonicolor is only asssociated with
Abies(alba). But this species has almost no bloom
of green and therefore it would be very surprising
if this species produces metallic green and gray
mycorrhizae as ACSAI & LARGENT mentioned. The
plectenchymatic mantle (prosenchymatic in the
authors’ terms) apparently resembles the mantle of
L. deterrimus. Unfortunately further descriptions
of the mycorrhizae by ACSAI & LARGENT are not
provided.
MELIN (1924) used an orange milking Lactarius from
gymnosperm forests (determined as L. deliciosus)
for his successful syntheses of mycorrhizae with
Pinus montana. He showed a cross section of the
mycorrhiza. A dense plectenchymatous mantle - as
we found for L. deterrimus - and a Hartig net with
several rows of hyphal cells are the most
important features shown by MELIN. He remarks that
the emanating hyphae break off very easily, as we
found for the mycorrhiza of L. deterrimus on Picea
abies.
HATCH & DOAK (1933) showed, for the purpose of
explaining some features of root systems the habit
and a longitudinal section of the synthesized
mycorrhiza of Lactarius deliciosus + Pinus strobus
but they gave no detailed description. Their
photographs of rhizomorphs of the synthesized
mycorrhizae are noteworthy.
It is very surprising that none of the above
mentioned authors have found laticiferous hyphae.
ZAK (1971a) described only the morphology, colour
and natural changes of colour of synthesized
mycorrhizae of Lactarius sanquiflvuus + Pseudotsuga
menziesii.
as)
Ze HACTARLUS, PLOLINUS Fr.
Pee) oS 6 Pl 345
MORPHOLOGICAL CHARACTERS
Ramification: monopodially pyramidal; length of
unramified ends: - 0,4 (0,5) mm; diameter of
unramified ends: (0,35) 0,42 - 0,45 (0,6) mm;
shape of unramified ends: straight, seldom
slightly bent; structure of surface: more or less
smooth, with some rhizomorphs.
Colour of unramified ends: fleshy-brown with
white, farinose cover; colour of tips: like the
other parts; colour of older. parts:) light
umbre-brown.
ANATOMICAL CHARACTERS OF SURFACE
OUTER SURFACE OF MANTLE
Structures plectenchymatous with short cylindrical
or ovoid projecting hyphal ends, intermixed with
Slightly yellowish cells with tiny granular
contents, laticiferous hyphae present, 7 - 9 pm in
diam., slightly yellowish, walls up to 1 pm thick;
size of hyphal cells: 3 - 4 pm in diam.; thickness
of cell walls: inconspicuous; surface of the very
tip: slightly pseudoparenchymatous, without or
only occasionally with short cylindrical or ovoid
projecting hyphal ends.
RHI ZOMORPHS
Shape in cross-section: more or less round;
arrangement and diameter of hyphae: central hyphae
more or less parallel to the axis, the innermost
10 - 15 (20) pm thick, remaining central hyphae
and irregularly woven ones, and projecting hyphae
3 - 4 pm thick; thickness of cell walls: central,
thick hyphae up to 2,5 pm, remaining hyphae 1
(1,5) pm; colour of hyphae: more or less
colourless; septae: clampless; anastomoses: rare,
hyphae mostly connected by strong agglutination.
EMANATING HYPHAE
Shape and diameter: occasionally ramified, with
16
blunt ends, ca. 2 pm in diam.; thickness of cell
walls: slightly thick-walled, thinner than 0,5 pm;
colour of hyphae: more or sets colourless; septae:
clampless, rare.
CYSTIDIA none, exclusive of the special,
projecting hyphae of the surface.
INNER SURFACE OF MANTLE
Arrangement of hyphae: mostly irregular, but
sometimes with strandlike arrangements, short but
more often relatively long; diameter of hyphae:
Zp ay VO) Bile
ANATOMICAL CHARACTERS, CROSS-SECTION
MANTLE
Thickness and differentiation: 40 - 60 po,
distally thicker than on the remaining parts of
the mycorrhizae, differentiated into three layers;
cell-shape and hyphal dimensions of different
layers: outermost layer with radially arranged
hyphal ends, radially 3 - 10 (15) pm, tangentially
3 - 6 (10) pm, innermost layer tangentially 5 - 10
um, “radially"3) =— 5 pm, remaining layer with hyphae
2 - 3,5 pm in diam, slightly tangentially longish,
laticiferous hyphae 4,5 - 8 (10) pm thick.
TANNIN CELLS
Number of rows: 1 - 2; shape and dimensions:
narrow or broader elliptic, outer row slightly
narrower than inner, 30 - 50 (80) x (5) 7 - 10
(20) pm.
HARTIG NET
Shape and dimension of cortex cells: round or
slightly elliptical, mostly radially oriented,
nearest to the endodermis often tangentially
oriented, (12) 15 - 25 (30) pm; depth of H'net: 4
- 5 rows inclusive of tannin cells; thickness of
H'net: between tannin cells 2 - 5 pm, conspicuous,
between cortex cells 2 - 3 m, inconspicuous;
number of hyphal rows of H'net: mostly 1, between
tannin cells often 2, outer tannin cells ‘mostly
Fig. 4 - Lactarius picinus - a: Habit of
mycorrhizae. - b, b', b'': Different surface
views. - c: Surface view of mycorrhiza, densely
covered with soil particles. - d: Surface view of
the very tip. - e: Inner surface of mantle. (All
figs. from RA 10745 in Herb. R. Agerer).
i]
1
18
separated by triangular projections of the inner
mantle layer; shape of H'net cells in section:
between and near tannin cells more or less round,
between cortex cells probably mostly radially
elongated, cell walls inconspicuous.
ANATOMICAL CHARACTERS, LONGITUDINAL SECTION
MANTLE
Shape _ and dimensions of the hyphae of the very
tips mantle 40 - 60 pm thick, only two layers
present, innermost layer not specialized, middle
layer distinctly thicker, hyphae more or less
parallel to the surface of root, hyphae of this
layer somewhat thicker than those of the remaining
partvyor Che mycorrnizayni 5.73 om?
differentiation through older parts of the mantle:
mantle only 35 - 45 um thick, with three layers;
shape and dimensions of cells in different layers:
outermost layer with hyphae tangentially 3 - 6
(10) pm, jj radially (3)—10"(15)'.nm,; 7 innermostwia ver
with hyphae tangentially (3) 5 - 10°(15)) pa,
radially (2)''3) -\6 pm, remaining layer with hyphae
tangentially (3) Sy o. 420) pm, radially aaa
(3) pm, laticiferous hyphae tangentially 5 - 30
(45) Digit aG Meili i 50 wos.
HARTIG NET
Shape and dimensions of tannin cells: irregularly
cylindrical or irreqularly longish, parallel to
the axis or oriented ina slightly oblique way to
inside backwards, (30) 60 - 100 (120) x 10 - 20
(30) pm; shape and dimensions of cortex cells:
irregularly rectanqular or irregularly rhombic,
oriented ina slight or conspicuous oblique way to
inside backwards, 40 - 80 (100) x (15) 20 - 25
(30) pm.
Fiq. 5 - Lactarius picinus - a: Longitudinal
section through rhizomorph. - b: Longitudinal
section through rhizomorph, detail from the
centre. - c: Surface view of laticiferous hyphae.
Arrows indicate the distal direction. (All fidqs.
from RA 10745 in Herb. R. Agerer).
19
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20
COLOUR REACTION IN DIFFERENT REAGENTS
Acetic-Fuchsin: slightl ink, laticiferous hyphae
artly intensivel ink; a-Naphtol: ----;
a
Ammonium-Hydroxide: ----; Brilliant-Kresyl-Blue:
blueish-green, laticiferous hyphae qreen, after
treatment with water violet blue, laticiferous
hyphae distinctly bluish-green; Chlorazol-Black Es:
qareyish-blue, laticiferous hyphae more intensive;
Cotton-Blue: bluish; Erythrosin: pinkish-brown,
laticiferous hyphae plasmatically red; Fast-Green:
reen, laticiferous hyphae dark green; Gujak:
; Lron-Sulfate: greenish-brown; KOH 15%: ----;
Fig. 6 - Lactarius picinus - a: Emanating hyphae
from mycorrhiza. - bs: Emanating hyphae from
rhizomorph. (All figs. from RA 10745 in Herb. R.
Agerer).
ai
Kongo-Red: brownish-red; Maqnesium-Acetate: ----;
Malachit-Green: green, laticiferous hyphae more
intensive; Martius-Yellows slightly yellowish,
laticiferous hyphae a little more intensive;
Melzer’s Reagent: yellowish-brown, laticiferous
hyphae distinctly brownish; Phenol: ----;
Phenol-Anilins ----; Phloxin B: pinkish-brown,
laticiferous hyphae more intensive; Stahl's
EP-Reagent: ----3; Sulfo-Vanillin: light red,
laticiferous hyphae dark carmine; Toluidin-Blue:
violet blue,
AUTOFLUORESCENCE
WHOLE MYKORRHIZA
UV 254 nm: ----; UV 366 nm: ----.
MANTLE AND RHIZOMORPHS
UV-filter 340 - 380 nm: laticiferous hyphae
distinctly blue-green, remaining hyphae slightly
blue-green; BLUE-filter 450 - 490 nm: laticiferous
hyphae distinctly green remaining hyphae slightly
green.
STAINING OF NUCLEI
Cells of projecting hyphae and longer hyphae below
these cells mostly with 2 nuclei, lying close
together, longer hyphae seldom with more than two
nulei ( 4 - 6). (Method: aceto-carmine).
Siderophilous reaction negative.
MATERIAL STUDIED AND METHOD OF IDENTIFICATION
Germany, Bavaria, Buchenau bei Flirstenfeldbruck,
2. 10. 1984; with Picea abies; reference specimen
or mycorrhiza and fruithpody RA 10745.in Herb. R.
Agerer. There were some connections between base
of fruitbody and mycorrhizae by rhizomorphs.
COMPARISON OF THE MYCORRHIZA OF LACTARIUS PICINUS
WITH THOSE OF L. DETERRIMUS
(See comments on L. deterrimus)
Le
23
MYCORRHIZAE OF LACTARIUS PICINUS AS TREATED IN
LITERATURE
No mycorrhiza formed by this species has been
discribed. This is true for all species of the
section Plinthogali of this genus.
Plate 4: overleaf
Pl. 4 - Lactarius picinus - a: Inner surface of
mantle. - b: Cross section. - c: Longitudinal
section through the very tip. - ds: Cross section.
(All figs. from RA 10745 in Herb. R. Agerer). -
Ban, for allyfigs. 10 \pm.
Pl. 3 - Lactarius picinus - a: Habit of
mycorrhizae. - b: Autofluorescence of laticiferous
hyphae (Blue-filter). - c: Rhizomorph,
longitudinal section. - d: Surface view of
laticiferous hyphae. - e: Rhizomorph, cross
section. - f: Inner surface of mantle. (All figs.
from RA 10745 in Herb. R. Agerer). - Bar 1 mm
fag. a), 10 pm (figs. b - £).
24
NN, -
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Hy
ating
Z5
3. RUSSULA OCHROLEUCA (Pers.) Fr.
Palate), pulp ne 26 aD 9 a Ole
MORPHOLOGICAL CHARACTERS
Ramification: monopodial; length of unramified
ends: - 2,8 (3,2) mm; diameter of unramified ends:
0,33 - 0,46 mm; shape of unramified ends: more or
less straight or only slightly bent; structure of
surface: smooth but with dots and with some
rhizomorphs.
Colour of unramified ends: yellowish or slightly
yellowish-olive with striking, yellowish dots of
small amorphous crystals; colour of tips: similar
as the other parts but without the yellowish dots
or with dots less conspicuous; colour of older
parts: umber brown with slightly olive tint,
yellowish dots not conspicuous or absent.
ANATOMICAL CHARACTERS OF SURFACE
OUTER SURFACE OF MANTLE
Structure: pseudoparenchymatous with angular cells
in surface view; size of hyphal cells: on the
average 8,3 cells cut in a square of 400 pm;
thickness of cell walls: up to 1 pm; surface of
the very tip: like the cells of the remaining
surface but cells smaller, about 17 cells cut ina
square of 400 pm.
RHIZOMORPHS
Shape _in cross-section: more or less round;
arrangement and diameter of hyphae: hyphae run
more or less parallel including a very thick
central hypha, central hypha 10 - 12 pm in diam.,
the other hyphae (1,5) 2 - 3 (4) pm in diam.;
thickness of cell walls: central hyphae up to 0,5
(1,0) pm, remaining hyphae inconspicuously
thickwalled; colour of hyphae: more or less
colourless; septae: without clamp connections;
anastomoses: rare, hyphae agglutinated.
EMANATING HYPHAE
Shape and diameter: (1,5) 2 - 3 pm in diam.;
26
Fig. 7 - Russula ochroleuca - a: Habit of
mycorrhizae. - b: Different surface views, with
granular patches of yellow pigment. - Cc: Different
surface views of the very tip. - ds: Inner surface
of mantle. (All figs. from RA 10705 in Herb. R.
Agerer).
pay
thickness ofscell walls: up to.0,5 pm; colour of
hyphae: more or less colourless; septae: without
clamp connections.
CYSTIDIA none found.
INNER SURFACE OF MANTLE
Arrangement of hyphae: irregular; diametre of
nyonaesi2) — 3 (4) um, .with short celis,or cells
somewhat longer.
ANATOMICAL CHARACTERS, CROSS-SECTION
MANTLE
Thickness and differentiation: 30 - 45 (50) pm,
pseudoparenchymatous throughout, with raised
spots; cell-shape and dimensions of different
layers: cells of the outermost layer furnished
partially with conical cell aggregates, mostly
filled with small granula, cells tangentially 10 -
Por 20) wie radrally — 3,5 =) 5S). pM, OUucermost
eo mi wan Oy) <0) pm thick, cellsvof)the innermost
fayer roundish, 2 -—- 3 (5) pm in diam., remaining
Pes cy Langentialivye 100-15 (20) jum, radially,+ 5 -
2 mM.
TANNIN CELLS
Number of rows: 1 - 2, the inner layer less
densely filled; shape and dimensions: flat and
compressed, the outermost row shorter than the
inner row, outer row - 30 - 55 (70) x 6 - 11 pn,
the innermost row - 20 - 35 (40) x 9 - 20 pm.
HARTIG NET
Shape and dimensions of cortex cells: round or
Slightly tangentially oval, 20 - 35 (60) x 15 - 25
fo) pm; depth of H'net: 4 -'5 rows) of cortex
cells; thickness of H'net: 2,5 - 3,5 (4) pm;
number of hyphal rows of H'net: 1; shape of H'net
cells in section: angular or slightly round,
quadratic or rectangular.
ANATOMICAL CHARACTERS, LONGITUDINAL SECTION
MANTLE
Shape and dimensions of the very tip: almost
regularly rectangular like a stone-wall, outermost
mayer 5 - 9.x 2,5 - 3 pm, innermost layer round to
quadratic ca. 2 pm in diam., remaining cells 5 - 9
29
x (2,5) 3 - 4 (5,5) pm, surface furnished with the
same conical cell-aggregates as mentioned for the
cross-section.
HARTIG NET
Shape and dimensions of cortex cells: flat-rhomic,
(40) 60 - 80 (100) x (15) 20 - 25 pm; shape and
dimensions of tannin cells: oriented in a oblique
way to inside backwards, (40) 50 - 80 (100) x (6)
See (016) um.
COLOUR REACTION IN DIFFERENT REAGENTS
Acetic-Fuchsin: salmon-red; a-Naphtol: ----;
Ammonium-Hydroxide: ----; Brilliant-Kresyl- _Blue:
bright blue after treatment with water violet
blue; Chlorazol-Black E: greyish-green;
Cotton-Blue: walls slightly blue; Erythrosin:
slightly pink; Fast-Green: yellowish-green; Gujak:
----; Iron-Sulfate: ----; KOH 15%: conspicuously
ochre; Kongo-Red: slightly orange;
Magnesium-Acetate: ---; Malachit-Green: greenish;
Martius-Yellow: ---; Melzer's Reagent: ---;
Phenol: dirty yellowish- geen; Phenol- eeuteiar ----;
Phloxin B: ---; Stahl's EP-Reagent: slightly
ochre; Sulfo-Vanillins: ----; Toluidin-Blue: violet
blue .
AUTOFLUORESCENCE
WHOLE MYKORRHIZA
UV 254 nm: ----; UV 366 nm: ----.
MANTLE AND RHIZOMORPHS
UV-filter 340 - 380 nm: slightly yellowish-qreen;
BLUE-filter 450 - 490 nm: distinctly
yellowish-qreen.
SSS
Fig. 8 - Russula ochroleuca - a: Rhizomorph in
surface view. - a': Squashed rhizomorph with free
central, thick hypha. - a'': Ramified central
hypha of rhizomorph. - b: Emanating hyphae of
mycorrhiza. - c, d: Bases of emanating hyphae, c
in tangential view, d in longitudinal section.
(All figs. from RA 10705 in Herb. R. Agerer).
30
STAINING OF NUCLEI
Cells of mantle and emanating hyphae with two
nuclei, lying close together (Method:
aceto-carmine).
Siderophilous reaction negative.
MATERIAL STUDIED AND METHOD OF IDENTIFICATION
Germany, Bavaria, zwischen Odelzhausen und Mering,
im Hédglwald, 19. 9. 1984, with Picea abies;
reference specimen of mycorrhiza and fruitbody in
Herb. RA 10705 in Herb. R. Agerer. There were
connections between the stipe of the fruitbody and
the mycorrhiza.
COMPARISON OF THE MYCORRHIZA OF RUSSULA OCHROLEUCA
WITH THOSE OF R. XERAMPELINA
(Selected features only)
(1) Morphological characters
A very conspicuous difference between these
two species concerns the colour of the
mycorrhizae. The mycorrhizae of Russula ochroleuca
are always furnished with striking yellowish dots
made up of conical cell-aggregates and granular
substances, which may lay upon the cell walls;
very often the cells of the conical
cell-aggregates are filled with qranules which may
cause the yellow colour, too. Similar encrusting
pigments are present on the hyphae of the cap
surface of fruitbodies. This is the reason why the
colour of the mycorrhiza is similar to the colour
ER
Pl. 5 - Russula ochroleuca - a: Habit of
mycorrhizae. - b: Autofluorescence of mantle
surface (Blue-filter). - c, e: Inner surface of
mantle. - d: Surface view of mantle. (All figs.
from RA 10705 in Herb. R. Agerer).. - Bar’ 1) mm
(fig. a), 10 pm (figs. b - e).
ui
a
32
of the fruitbody. The same is recognizable in R.
xerampelina, though there are obviously no
encrusting pigments on the mantel surface, the
dark fleshy-brown appearence is reminiscent of the
less intensively coloured parts of the fruitbody
of R. xerampelina,
The most striking differences occur once more
in the structure of mantles. The mantle surface of
R. ochroleuca shows a typical pseudoparenchyma
with angular cells furnished with the conical
cell-aggregates, whereas R. xerampelina possesses
a more or less dense network of strongly
anastomosing hyphae. Only the mantle of R.
xerampelina shows typical cystidia as known in the
genus Russula (ROMAGNESI 1967). R. ochroleuca
seems, to’ produce thicker mantles) than yaa
xerampelina . The rhizomorphs, however, belong to
the same type with centrally arranged voluminous
hyphae. It is probably not of great significance
that there are a lot of such thick hyphacein sk.
xerampelina but only a single one or a few in Re
ochroleuca. The central hyphae of the rhizomorphs
of R. ochroleuca seem to be thicker than those of
R. xerampelina.
(2) Characterization after treatment with
different reagents
Many differences could be found, but some are
not distinctive enough to differentiate the two
species. The most conspicuous are the following.
Chlorazol-black E makes the mantle of R.
ochroleuca mycorrhizae qreyish-green whereas it
turns the mantle of R. xerampelina greyish-blue.
The mantle of R. ochroleuca becomes distinctly
ochraceous after treatment with KOH whereas the
mycorrhiza of R. xerampelina does not show this
Pl. 6 - Russula ochroleuca - a, b: Emanating
hyphae (longitutinal section). - c: Cross section.
- d: Longitudinal section, somewhat more distant
from very tip than fiq. f. - e: Longitudinal
section through very tip. - f: Longitudinal
section, behind the very tip. - g: Longitudinal
section, mantle with conical cell aggregates. (All
figqs. from RA 10705 in Herb. R. Agerer). - Bar for
ali figs, 10 pm.
i
ah
ss We
34
reaction.) In Phenol) the mycorrhiza of KR.
ochroleuca becomes dirty yellowish-green, while
that of R. xerampelina shows no reaction. A
further difference regarding treatment with Phenol
is that after washing of the tinged mantle with
water the mantle of R. xerampelina alone becomes
slightly umber-brown. Phloxin B causes the mantle
of the R. xerampelina mycorrhiza to show a
slightly red reaction. Stahl's EP reagent,
however, turns the mantle of R. ochroleuca
exclusively into a slighty ochre colour. Only in
R. xerampelina were the inner hyphae of mantle
tinged distinctly red with Sulfo-Vanillin. In R.
xerampelina ROMAGNESI (1967) remarks that there is
no reaction with Sulfo-Vanillin. Only
insignificant differences could be recognized
after treatment with Cotton-blue: the hyphae of R.
xerampelina become darker than those of R.
ochroleuca. ROMAGNESI (1967) obtained a purplish
colour after having treated the yellowish
encrusted hyphae of the cap of R. ochroeuca with
acetic-fuchsin, a salmon red tinge was produced in
our ivestigations of the mycorrhiza of R.
ochroleuca. The mantle of R. xerampelina, however,
showed a similar reaction with acetic-fuchsin to
that of R. ochroleuca. SCHAFFER (1952) mentioned
an olive reaction of the fruitbodies of R.
xerampelina with iron-sulfate, which did not
correspond with our results on the mycorrhizae of
this species. According to SCHAFFER Russula
ochroleuca shows an indish-red reaction wih
Phenol, but the mantle of the mycorrhizae became
dirty yellowish-green with this reagent. These
differences between the reactions of fruitbodies
and mycorrhizae may well be caused by storage of
the mycorrhizae.
(3) Siderophilous granula
Russula xerampelina is so far the only
species of Russula with siderophilous reactions of
parts of the mycorrhizal mantle. No such reaction
is known in parts of the fruitbodies.
MYCORRHIZA OF RUSSULA OCHROLEUCA AS TREATED IN
LITERATURE
There is not a single mycorrhiza known of the
species of the section Ingratae of the genus :
Russula. |
he
4, RUSSULA XERAMPELINA (Schff. ex Secr.) Fr.
POs ey REL eh ph Oe
MORPHOLOGICAL CHARACTERS
Ramification: monopodial; length of unramified
ends: up to 2 mm; diameter of unramified ends:
(0,24) 0,33 - 0,39 (0,48) mm; shape of unramified
ends: more or less straight; structure of surface:
smooth, with some rhizomorphs.
Colour of unramified ends: dark flesh-brown,
lustreless; colour of tips: somewhat
paler,lustrous; colour of older parts; dark
reddish brown, lustrous.
ANATOMICAL CHARACTERS OF SURFACE
OUTER SURFACE OF MANTLE
Structure: losely woven, like a net of strongly
anastomosing hyphae; size of hyphal cells: (4) 5 -
fom, caeckness of icelliwaliss up Coul umes ysurtace
of the very tip: hyphae conspicuous, short,
densely plectenchymatous or slightly
pseudoparenchymatous.
RHIZOMORPHS
Shape in cross-section: more or less round;
arrangement and diameter of hyphae: central hyphae
Paick, wii 2 sum, remaining hyphae (2,75 - 5 pm;
thickness of cell walls: walls of central hyphae
ca. 1 pm, joining hyphae only sligtly thick
walled, hyphae towards the margin up to (0,5) 1
bm; colour of hyphae; more or less colourless,
septae: clampless; anastomoses: rare.
EMANATING HYPHAE
Shape and diameter: probably only proximally
ramified, 1,5 - 2 pm; thickness of cell walls:
only slightly thick-walled, thinner than 0,3 pm;
colour of hyphae: more or less colourless; septae:
clampless.
CYSTIDIA rare, somewhat subulate, 30 - 40 x 5 - 6
pm (proximal).
INNER SURFACE OF MANTLE
Arrangement of hyphae: irreqular, sometimes
36
coiled, hyphae densely woven, agqlutinated;
Gdiametre of hyphaes 2,5 - 3 pm.
ANATOMICAL CHARACTERS, CROSS-SECTION
MANTLE
Thickness and differentiation: 20 - 30 pm,
pseudoparenchymatous throughout; cell-shape and
dimensions of different layers: outermost hyphal
layer losely woven, tangentially (2) 6 - 10 (14)
pm, radially 2 - 2,5 pm, only slightly
thick-walled, innermost layer with more or less
round cells, 2 - 3 pm in diam., remaining cells
more or less round or slightly tangentially
elongated, tangentially (2) 5 - 9 (15) pm,
radially (2) 3 - 4 (5) jam.
TANNIN CELLS
Number of rows: 2 (-3); shape and dimensions:
irregularly shaped, tangentially-slender,
outermost row 50 - 65 x 2 - 87 (20) pm, joining
row (35) 45 = 60 (70) x 8 — 15 mim.
HARTIG NET
Shape and dimensions of cortex cells: oval,
slightly radially elongated, (20) 27 - 35 x 10 -
20 pm; deepness of H'net: 4 - 5 rows neighbouring
to the endodermis; thickness of H'net: 3 - 4 pm;
number of hyphal rows of H'net: 1 (between tannin
cells up to 2 (3) rows); shape of H'net cells in
section: often elongated, but cells of the outer
cortex layer mostly more or less round.
ANATOMICAL CHARACTERS, LONGITUDINAL SECTION
MANTLE
Shape and dimension of the very tip:
pseudoparenchymatous, with smaller cells, without
distinct layers, tangentially 3 - 5 pm, radially
Fig. 9 - Russula xerampelina - a: Habit of
mycorrhizae. - b, b', b'': Different surface
views. - c, c': Different views of inner mantle
surface. - d, da': Different views of the very tip.
(All figs. from RA 10725 in Herb. R. Agerer).
es :
i>
38
2 - 3 pm; shape and dimensions of cells in
different layers: outermost cells tangentially 6 -
10 (15) pm, radially 2 - 4 pm, innermost layer
tangentially 3 - 8 x 2 - 3 pm, remaining cells 5 -
TBA) Sai ak On) A'S Pon LIT s
HARTIG NET
Shape and dimensions of cortex cells: elongated,
oriented in an oblique way to inside backwards,
(30) 40° -60 (80) x (12) 15 - 22..(28) pm; shape’ and
dimensions of tannin cells: elongated like cortex
cells 30) 601 (0 vane 1 Omnis
COLOUR REACTION IN DIFFERENT REAGENTS
Acetic-Fuchsin: slightly reddish; a-Naphtol: ----;
Ammonium-Hydroxide: ----; Brilliant-Kresyl-Blue:
bright blue, after treatment with water violet
blue; Chlorazol-Black E: qreyish-blue;
Cotton-Blue: hyphal walls distinctly blue;
Erythrosin: slightly red; Fast-Green: green;
Gujak: ----; Iron-Sulfate: ----; KOH 15%: ----;
Kongo-Red: slightly red; Magnesium-Acetate: ----;
Malachit-Green: green; Martius-Yellow: ----;
Melzer's Reagent: ----; Phenol: _----, but after
treatment with water slightly umbre-brown;
Phenol-Anilin: ----; Phloxin B: slightly red;
Stahl's EP-Reagent: ----; Sulfo-Vanillin: hyphae
of inner surface of mantle distinctly pink, the
other hyphae only pinkish; Toluidin-Blue: violet
blue.
AUTOFLUORESCENCE
WHOLE MYKORRHIZA
UV 254 nm: ----; UV 366 nm: ----.
Fig. 10 - Russula_ xerampelina - as: Optical section
through rhizomorph running along rootlet. - b:
Emanating hyphae of mycorrhiza. - c: Emanating
hyphae (cross section of mycorrhiza). - d, d':
Longitudinal section (mantle). - e: Hartiq net
with balloon like structure (cross section). (All
figs. from RA 10725 in Herb. R. Aqerer).
40
MANTLE AND RHIZOMORPHS
UV-filter 340 - 380 nm: slightly blue; BLUE-filter
450 - 490 nm: distinctly qreenish-yellow.
STAINING OF NUCLEI
Cells of the inner mantle hyphae, of emanating
hyphae and cells of the hyphal net on the outer
surface had two nuclei lying mostly close together
(method: acetic-carmine). - The siderophilous
reaction was positive in some of the outermost
hyphae of the mycorrhizal surface.
MATERIAL STUDIED AND METHOD OF IDENTIFICATION
Germany, Bavaria, Munchen-Oberschlei&heim,
Korbiniani-Hd6lzl, bei Versuchsflache O, 22. 9.
1984, reference specimen of mycorrhiza and
fruitbody RA 10725 in Herb. R. Agerer; with Picea
abies. The Mykorrhiza was overgrown by the stipe
ofthe fruvtbody.
COMPARISON OF THE MYCORRHIZAE OF RUSSULA
XERAMPELINA WITH THOSE OF R. OCHROLEUCA
(See comments on R. ochroleuca)
MYCORRHIZAE OF RUSSULA XERAMPELINA AS TREATED IN
LITERATURE
Out of the section Rigidae of the genus Russula
only the mycorrhizae of Russula_vesca and R.
grisea are known. MODESS (1941) tried to obtain R.
xerampelina in culture but he was not successful.
Pl. 7 - Russula xerampelina - a: Habit of
mycorrhizae. - b, d: Inner surface of mantle. - c:
Autofluorescence of inner layers of mantle. -e -
g: Surface views in three different focus' from
one and the same point: e = outermost, f = joining
layer, g = a little more deeper. (All fiaqs. from
RA 10725 in Herb. R. Agerer). - Bar 1 mm (fig. a),
TOnine CfA gseib—1ig)).
a Ze
BA
\\
IN
\ \
\ Ta \
Cod,
i
\\
i
\ Ay
\Y
Ih ON
\
41
42
CERUTI & BUSSETTI (1962) described and
illustrated with half schematic line drawings the
mycorrhiza of R. qrisea + Tilia tomentosa. A
striking feature are the pear-shaped cystidia and
a dense indument of emanating hyphae on the mantle
surface. The brightly coloured sulfur-yellow
mycorrhiza and the light yellow colour of the
mantle distinguish this mycorrhiza very well from
that of R. xerampelina, too. The structure of the
mantle should be prosenchymatic. Further essential
characteristics are not mentioned.
LUPPI & GAUTERO (1967) described and drew what
is probably the mycorrhiza of R. vesca. The mantle
is pseudoparenchymatous at some points
plectenchymatous at others. In this feature the
mycorrhiza of R. cf. vesca seems to be similar to
that of R. xerampelina. The authors, however,
showed no surface view of the mantle and the
Similarity is therefore not obvious. The colour of
the mycorrhiza characterized by LUPPI & GAUTERO is
cream or light hazel-brown and thus deviates from
that of the mycorrhiza of R. xerampelina. The
emanating hyphae of these two species look
Similar.
SS SS
Pl. 8 - Russula xerampelina - a: Longitudinal
section through the very tip. - bs: Longitudinal
section. - c: Cross section near the very tip. -
ds: Cross section. (All figs. from RA 10725 ‘in
Herb. R. Agerer). = Bar for all itigs. 10. um.
43
a4
LLX DISCUSSION
1. Discussion of the features used for
characterization
a. Surface views (inner and outer) of mantle
This feature was first used by GIBELLI
(1883) and FRANK (1885). GIBELLI characterized
some mycorrhizae by line drawings of their
surface. His studies show remarkable details of
structure. He studied mycorrhizae of different
host trees: Quercus, Corylus,.Ostrya, Carpinus.
FRANK (1885) described mycorrhizae of Carpinus
betulus and of Fagus silvatica. He showed three
characteristic surface views: one showing a
plectenchymatous structure with many emanating
hyphae, another a plectenchymatous view with few
emanating hyphae, and a third of a smooth
mycorrhiza with pseudoparenchymatous surface and
without emanating hyphae. MANGIN (1910) also used
the surface view characteristics when he described
the mycorrhiza of Fagus silvatica and showed a
line drawing of it. Unfortunately the feature of
surface view of ectomycorrhizae was forgotten or
not taken into account until CHILVERS (1968) used
it for the description of Eucalyptus mycorrhizae.
He showed remarkable differences in this
characteristic and documented this with exact line
drawings. Because of the consequent usage of the
surface views the study by CHILVERS is one of the
most important and most instructive publications
concerning characterization of ectomycorrhizae.
ZAK (1969) used the surface view to characterize
the mycorrhiza of Poria terrestris + Pseudotsuga
menziesii. However, for characterizing the
mycorrhiza of Rhizopogon vinicolor + Pseudotsuga
menziesii ZAK (1971) and for Byssoporia ia terrestris
+ Pseudotsuga menziesii (ZAK & LARSEN 1978) ZAK
renounced the use of this feature with. VOIRY
(1981) too, did not show mantle surface views. He,
however, used tangential sections through the
mycorrhiza as did some other investigators. Such
sections may show some important features too, but
they are mostly not as characteristic as are
direct views of the surface (this is well
demonstrated by the photographs of different
levels of focus of the mycorrhizae of Russula
45
xerampelina, see plt. 7). ALEXANDER (1981) again
used this feature for characterizing the
mycorrhiza of Lactarius rufus + Picea sitchensis;
he applied an interference contrast microscope to
demonstrate the epidermoid structure of this
smooth mycorrhiza. Thus he was able to get a more
impressive view. PALM & STEWART (1984) depicted
the surface view of only one of their
characterized mycorrhizae of some Suillus species.
Even Godbout & Fortin (1985a) did no show the
surface views of the synthesized mycorrhizae with
Populus tremuloides and they "felt that this is
an informaton gap. In future, however, they will
use tangential views and rhizomorphal structures
as CHILVERS (1968) has done.
Currently, scanning electron microscope is more
often used. But it seems to be of limited value,
because it does not show advanced features. And a
SEM is unsuitable for determining of mycorrhizae.
In addition light microscopy shows features of
septation of hyphae. In relation to the structure
of crystals which may sheath the hyphae, however,
SEM may be very useful (THOMAS & JACKSON 1982).
The same is true for ontogenetic studies (PICHE et
al. 1983). The development of the mantle was
researched by SEM very successfully for example by
SrRuUgcLU (1973, 1979) and SEVIOUR et al. (1978).
It appears that ZAK (1969) and PALM & STEWART
(1984) are the only who studied and described the
inner surface of the mycorrhizal mantle. ZAK was
able to demonstrate that this view may show some
instructive details. The innermost layer of
Byssoporia terrestris + Pseudotsuga menziesii
possessed stellate arrangement of hyphae. In our
researches on further mycorrhizae we found similar
structures (AGERER, in preparation).
Another advantage of surface view is that it
enables examination of the structure of
laticiferous hyphae of mycorrhizae of deeper
layers of the mantle. Such features were used
successfully by LUPPI & GAUTERO (1967) and VOIRY
(1981).
b. Rhizomorphs
In the literature several terms have been
employed to describe these structures. For
example, rhizomorph, hyphal bundle, and hyphal
46
strand. The term rhizomorph is mostly used for
compactly packed hyphae running more or less
parallel whereas hyphal bundle or strand is used
if hyphae are more or less loosely connected by
different mechanisms, e. gq. anastomoses or
twirling around. Because a continuum can be
demonstrated (AGERER, in prep.) between the two
extremes we use exclusively the term rhizomorph
when hyphae are connected and grow more or less
parallel and more or less close together for a
greater distance.
Again it was CHILVERS (1968) who used the
features of the rhizomorphs consequently to
describe a number of mycorrhizae. The variety of
organization of rhizomorphs is impressive. Besides
the mantle view this structure may help a great
deal to differentiate between mycorrhizal types,
as for example inthe case of mycorrhizae of
Lactarius deterrimus, L. picinus, Russula
ochroleuca and R. xerampelina.
The rhizomorphs are likely to be
physiologically important for the so-called smooth
mycorrhizae known especially within the genera
Lactarius and Russula (FROIDEVAUX 1973, LUPPI &
GAUTERO 1967, MELIN 1924, OGAWA 1981, RIFFLE 1973,
VOIRY 1981). Because usually very few emanating
hyphae are present, the transport of water must
occur principally within the rhizomorphs. There
exist different types of rhizomorphs even in one
genus (see Lactarius deterrimus and L. picinus).
Our results do not concur with those of GODBOUT &
FORTIN (1985) who characterized the mycorrhizae of
Lactarius and Russla as being without strands
(rhizomorphs). All mycorrhizae of these two genera
which we have examined so far (AGERER, in prep.
BRAND & AGERER in prep., GRONBACH & AGERER in
prep.) showed typical rhizomorphs.
It seems that where rhizomorphs could not be
found they may have been lost in the extraction |
and cleaning process. It is unlikely that the
sparse emanating hyphae are effective enough to
nourish the mycorrhizae and the tree. Comparing
mycorrhizae from soil and those grown by t
synthesizing ALEXANDER (1981) remarked for
Lactarius rufus that in nature he had never found
rhizomorphs whereas they appeared on the i
synthesized mycorrhizae. The supposition that
47
rhizomorphs are probably present and lost only by
washing is supported by AGERER (1985) who made it
probable that the mycorrhizae of the genera
Lactarius and Russula belong to the most effective
ones for water transport. And this would not be
understandable for mycorrhizae without rhizomorphs
or without enough effective emanating hyphae.
MASUI (1926a) stated for an unidentified
mycorrhiza that old ones in contrast to young
mycorrhizae always produce hyphal bundles or
filaments. Therefore one must additionally take
into account that the presence or absence of
rhizomorphs could be depending on the age of
mycorrhizae.
Features of the rhizomorphs were used for
delimiting types of mycorrhizae by VOIRY (1981),
but unfortunately no details of the rhizomorph
structures were given.
In the early twenties MASUI (1926b)
observed hyphal bundles on a Quercus mycorrhiza
and showed a cross section through a rhizomorph
which he believed belonged to Suillus luteus. For
the fungs he demonstrated thick centrally arranged
hyphae. In the same year (MASUI 1926a) MASUI
described and showed with line drawings some
rhizomorphs of different mycorrhizae. They were
partially differentiated into centrally thicker
hyphae and diverging thinner ones. MASUI (1926,
1927) figured the morphology of some smooth
mycorrhizae possessing rhizomorphs without
characterizing them microscopically. MELIN (1923)
provides line drawings of a rhizomorph with
distinctly thick, voluminous central hyphae
surrounded by thin parallelly running hyphae,
otherwise he only described or depicted the gross
morphology of rhizomorphs.
ZAK (1969) compared microscopically the
rhizomorphs of Byssoporia terrestris (var.
sartoryi and var. aurantiaca) and was able to show
distinct differences in the organization of the
hyphae. One rhizomorph possessed more or less
parallel hyphae whereas the other was built of
irregularly interwoven hyphae. The rhizomorphs of
Rhizopogon vinicolor + Pseudotsuga menziesii
mycorrhizae were also described anatomically by
ZAK (1971). In comparison with those of B.
terrestris there were distinct differences.
48
Rhizopogon vinicolor produced rhizomorphs with
centrally arranged thick hyphae. Again ZAK (ZAK &
LARSEN 1978) analyzed microscopically the
structure of rhizomorphs of some additional
varieties of Byssoporia terrestris. These
rhizomorphs showed striking differences from those
of the at first mentioned varieties (ZAK 1969) of
this species. These examples make it obvious that
the features of rhizomorphs are as useful as those
of the surface views for characterization of
ectomycorrhizae. The taxonomic importance of the
rhizomorphs was already stressed by ZAK (1973).
PALM & STEWART (1984) described the mycorrhizae of
some Suillus species and showed rhizomorphs.
Microscopical descriptions of the rhizomorphs,
however, are absent.
Recent investiqations on organization and
function of rhizomorphs were carried out by FOSTER
(1981) and DUDDRIDGE et al. (1980).
c. Cross- and longitudinal sections, emanating
hyphae and cystidia
These features were applied by DOMINIK
(1969) for characterizing various mycorrhizae and
for constructing a key. Since DOMINIK these
characteristics have very often been successfully
used. Organization of mantle (prosenchymatous,
pseudoparenchymatous), type of cystidia, features
of the emanating hyphae (colour, thickness of
walls, clamps, diameter a.s.o.) are all suitable
to describe mycorrhizae.
In our investigation only one type of
cystidia occurred (a typical one of the genus
Russula in R. xerampelina mycorrhiza), emanating
hyphae are very scanty.
Longitudinal sections have seldom been
investigated. LUPPI & GAUTERO (1967) described
different mycorrhizae showing only longitudinal
sections. Whereas MASUI (1926a, 1926b, 1927) and
CHILVERS (1968) applied cross- and longitudinal
sections for describing mycorrhizae. |
We were able to point out that longitudinal |
sections are important for characterizing the
mycorrhizae, because the structure of the mantle
may vary from the very tip to its proximal parts
(see R. ochroleuca, pl. 6).
49
d. Reactions with different chemicals
ZAK (1973) reviewed the publications which
have dealt with colour reactions of mycorrhizae
with different reagents up to 1973.
With the aid of chemical reagents FROIDEVAUX
(1973) compared the mycorrhiza of Lactarius
obscuratus with the basal mycelium of the
fruitbodies; he used eight different reagents
(Fron-suitate, Sulfuric’acid, KOH 15%;
Ammonium-hydroxide, Chlorovanillin, Guaiacol,
Pyrogallol, Sulfoformol) and he was able to show
corresponding colour changes.
MISHRA & SHARMA (1981) compared with the aid
of chemical reagents mycorrhizae with associated
fruitbodies. They used five different reagents
(Iron-sulfate, Sulfuric acid, Ammonium-hydroxide,
KOH (15%) and Melzer's reagent). They found on the
one hand differences between species of
mycorrhizae and on the other between mycorrhizae,
rhizomorphs and fruitbodies of one and the same
species.
In our present studies of ectomycorrhizae we
used a set of chemicals comprising now 22
different reagents. These chemicals are already
applied to about four dozen still undescribed
mycorrhizae. And it seems that some of these
reagents cause in all mycorrhizae the same colour
changes, for example fast-qreen and
malachit-qreen. It is possible that therefore some
of them will not be applied further. Other
reagents will be added. It is the aim to get a
suitable set of reaqents to characterize
mycorrhizae on a chemical basis.
e. Autofluorescence
TRAPPE (1967) and ZAK (1973) proposed to use
the autofluorescence of mycorrhizae at different
wave lengths. This approach was has been used
rather rarely although some mycologists use it for
characterizing relationships of fungi (MOSER 1978,
subgenus Leprocybe of genus Cortinarius).
Fluorescence microscopic has been used even less
often. ZAK (1971) studied the autofluorescence of
the whole mycorrhiza of Rhizopogon vinicolor,
FROIDEVAUX (1973) that of pieces of mantle of
Lactarius obscuratus mycorrhizae. The differences
50
revealed between the four mycorrhizae in the
present study using this method are not exciting,
but further studies (AGERER, in prep.) have
already shown that there are striking differences
in the behaviour regarding autofluorescence.
f. Staining of nuclei and siderophilous
granulation.
Up to now the siderophilous granulation has
not been used for characterization of mycorrhizae
but this characteristic has been widely applied by
fruitbody taxonomists (e.q. CLEMENCON 1978). As
has been shown, the mycorrhizae of Russula
xerampelina possess siderophilous granula in some
of the hyphae of the mantle surface; this
character will be studied in further
investiqations. CLEMENCON (1978) noted for Russula
Eragilis, RR. sardonia,and R. adusta)and) for
Lactarius maculatus and L. griseus small
siderophilous granula in the basidia; Russula
ochroleuca, R. xerampelina, Lactarius deterrimus
and L. picinus have probably not been studied yet
in this regard.
Nuclear staining reactions provide the
opportunity to distinguish between mycorrhizae
produced by ascomycetes and by clampless
basidiomycetes. All mycorrhizae without clamps
examined up to now have mostly had cells with two
nuclei. It may be important to know, whether the
nuclei lie close together or not (AGERER, in
prep.).
q. Shape and colour
ZAK (1973) reviewed the value of colour and
form in mycorrhizae. PALM & STEWART (1984) were
able to stress differences in the shape of Suillus
mycorrhizae. They could categorize the mycorrhizae
in terms of the mode of branching and the
arrangement of branches.
The shape of the presently treated
mycorrhizae is relatively uniform, and the
pyramidal shape of the mycorrhizae of Lactarius
picinus can be considered merely as a somewhat
denser ramification arising from a pinnate
arrangement. The colours of the characterized
mycorrhizae are typical and distinct. The
variability of colour is stressed by the
Dl
mycorrhiza of Lactarius deterrimus which changes
the colour with increasing age from bright copper
orange to dark olive. A fact which is described by
ZAK (1973) for the mycorrhiza of Lactarius
sanquifluus + Pseudotsugqa menziesii.
2. Methods of identification
Syntheses experiments have the disadvantage
that it is not possible to simulate natural
conditions for the fungi or for the tree. Some
publications make it obvious that there are
alterations of mycorrhizal types with ageing of
trees (e.q. CHU-CHOU 1979, FLEMING et al. 1984).
And therefore it could be quite possible that the
influence of the youth of tree-seedlings may be
great enough to change some features of the
mycorrhizae compared with those of mycorrhizae
known from old trees. There are some other known
effects on the structure of mycorrhizae. DUDDRIDGE
& READ (1984) have shown the influence of sugar
within the nutrient solution for the development
of mycorrhizae and were able to detect distinct
alterations in the area of mantle and Hartig net.
The influence of pH was already shown by MELIN
(1924a) and was stressed by MOSSE et. al. (1981).
The importance of nutritional factors was
demonstrated e. q. by MEYER (1985) very recently.
Some other factors may play an important role in
the formation of mycorrhizal structures such as
light intensity and age of the fungal inoculum.
Although MELIN (1959) stressed that the
mycorrhizae synthesized by him had the same
appearance and the same anatomical features
compared with those isolated form the soil,
detailed comparisons of synthesized and natural
mycorrhizae are very rare. Recently ALEXANDER
(1981) compared synthesized and natural
mycorrhizae of Lactarius rufus + Picea sitchensis
and discovered some differences in the mode of
ramification, in colour of mantle, in contents of
hyphae and in shape of the tannin-layer. Probably
these features are of less importance for
characterizing mycorrhizae, but it must be
considered that these may be morphological and
anatomical differences which may become essential
for delimiting species of mycorrhizae. The most
52
important characteristic - the shape of cells of
the mantle surface - however, remained quite
identical. The rhizomorphs could not be compared
by ALEXANDER because he could not find rhizomorphs
on mycorrhizae formed in natural soil.
PACHLEWSKI (1968) ascertained that with respect
to the method of synthesizing there may be
striking differences in the structure of
mycorrhizae. The mycorrhizae of Rhizopogon
luteolus, formed in the air, possessed a
distinctly hairy mantle with rhizomorphs (for
possible explanation see HARLEY & SMITH 1983),
whereas mycorrhizae grown in aqar were smooth or
only sporadically hairy. Differences in the cross
sections of the mantle were noted too, by
PACHLEWSKI. Hairy mantles were two-layered, while
smooth mantles were undifferentiated. Although the
differences between natural and aseptically grown
mycorrhizae may not always be as qreat as under
the extreme synthesizing conditions used by
PACHLEWSKI, these facts stress the need for
cautious interpretation of features seen in
synthesized mycorrhizae. THOMAS & JACKSON (1979)
also warn against relating aseptically synthesized
mycorrhizae with those occurring in nature. LAIHO
(1970), however, stated that in his researches on
Paxillus involutus there were no differences
recognizable between natural mycorrhizae and
mycorrhizae synthesized under different
conditions.
One of the most important questions is, how to
recognize mycorrhizae found in natural soil. A
decision about the identity of mycorrhizae is
really possible if we look critically and
patiently for unequivocal connections between
mycorrhizae and fruitbodies. This is especially
true if we can detect and follow distinct
rhizomorphs. For those mycorrhizal fungi which
cannot be cultured, this method is the one and
the only possibility to identify mycorrhizae with
certainty. If mycorrhizae were isolated from the
soil one can be shure that all characteristics are
natural in contrast to the features of the
synthesized mycorrhizae which well may provide
additional or changed features.
53
3. Variability of the species of mycorrhizae
Researches on the variability of
ectomycorrhizae isolated from soil have not so far
been concerned. This is not surprising, because
the descriptions of unidentified mycorrhizae are
mostly very poor and therefore a species concept
of such mycorrhizae could not be provided. The
same applies to identified mycorrhizae; partially
because of poor descriptions and partially because
identifications of mycorrhizae - by finding
connections between mycorrhizae and fruitbodies -
have been to few up to now. The same holds true
for the present investigation. Such studies of the
variability - inclusive of variability depending
on aqe - are urgently needed, but this must be the
task of time consuming monoqraphical studies. At
present it is a step forward to have single
specimens of mycorrhizae described precisely.
A further advance may be obtained by
studying the variability of ectomycorrhizae when
synthesized in different soils, under different
environmental circumstances and with a variety of
hosts. ALEXANDER (1981) attempted this approach
using the mycorrhizae of Lactarius rufus + Picea
Sitchensis. The synthesized mycorrhizae did not
show differences in response to different
conditions of syntheses. Such studies as well as
researches on the variation of mycorrhizae
isolated from soil would greatly assure in the
characterization and classification of
ectomycorrhizae,.
54
IV ACKNOWLEDGEMENTS
I am very much obliged to Mr. P. Altheide for having
skillfully prepared and stained sections and to Mr. K.
Liedl who has made the prints of the photos and drawings.
In addition I am thanking all members of my staff for kind
co-operation and discussing a lot of problems. Last not
least I am very thankful to Dr. David J. Read (Sheffield)
and Dr. Jim M. Trappe (Corvallis) for critical comments on
the manuscript and for improving the English text.
This study was supported by "DFG, Deutsche Forschungs
Gemeinschaft, grant Ag 7/5-2". Dachser Spedition (Munich),
Daimler-Benz AG (Stuttgart) and Carl Zeiss AG (Oberkochen)
susidized the investigations by a "fund for researching the
tree die-back".
55
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MYCOTAXON
VOue XV EL, apps? 61-62
October-December 1986
A NEW SPECIES OF THE GENUS OIDIUM FROM INDIA
G. Bagyanarayana
Osmania University, Department of Botany,
Hyderabad (A.P.), India
U. Braun
Padagogische Hochschule "Wolfgang Ratke”
Sektion Biologie/Chemie, WB. Botanik II
DDR-4370 Ko6then, Lohmannstr. 23, G.O.R.
Oidium pavoniae G. Bagyanarayana & U. Braun spec. nov.
Mycelio amphigeno, albo. Hyphis sterilibus rependi-
bus, hyalinis, ramosis, flexuosis, septatis, appresso-
riis non lobatis vel lobatis. Conidiophoris simplicibus,
ex hyphis sterilibus oriundis, cellulis ad basim cylin-
conidio- Sea,
30 wm
fs 5 «Bye e 00 + mee ot
Free Dae rei Biotic each oe
Ee CS Soke FR . .
} o se
cea Neate oy ee
Fig. 1. O. pavoniae spec.
nov.
draceis, rectis, 45-80 x 6.5-
10 um, conidiis solitariis,
ellipsoideis -cylindraceis vel
cylindraceis - doliiformibus,
(30-) 35-50 x 14-21.5 pum.
Holotypus: India, Hyderabad,
rayana; type host - Pavonia
zeylanica Cav. (HAL).
Mycelium amphigenous, white,
patches or effused, hyphae
hyaline, septate, branched,
ca 2.5-8 pm wide, appressoria
nipple-shaped to slightly
lobed, conidiophores erect,
foot-cells straight, cylin-
dric, long, 45-80 x 6.5=-10 um,
followed by 1-3 shorter cells,
the second cell sometimes as
long as the foot-cell, coni-
dia formed singly, ellipsoid -
cylindric or cylindric - doli-
form, (30-) 35-50 x 14-21.5
MN wtFigs. 1.
62
The known Oidium species on host plants of the Mal-
vaceae are distinguished from O. pavoniae by the follo-
wing features: Oidium abelmoschi Thum. (conidia in chains,
smaller), QO. schmiedeknechtii U. Braun (foot-cells of the
conidiophores short, ca 30-40 um long), O. urenae Yen
(foot-cells of the conidiophores very long, ca 60-200 pm,
conidia ovoid-doliform).
MYCOTAXON
Fs as ASP eI ee Oa sa a
Vol. XXVII, pp. 63-69 October-December 1986
CONIOCHAETIDIUM MIRABILE, A NEW ASCOMYCETE
ISOLATED FROM SALTED FOOD
Shun-ichi UDAGAWA! and Haruo TSUBOUCHI2
1 wational Institute of Hygienic Sciences, Kamiyoga
l-chome, Setagaya-ku, Tokyo 158, Japan
2 Nagoya City Health Research Institute, Hagiyama-
cho, Mizuho-ku, Nagoya 467, Japan
SUMMARY
A new species Of Coniochaetidium isolated from
salted food (dried powder of salmon and 'wakame'
(Undaria pinnatifida)) is described. The species,
C. mirabile, differs from Cc. ostreum, the type
species Of Coniochaetidium, in the morphological
characteristics in culture.
During a microbiological study of retailed salted foods
in Nagoya City of central Japan, the junior author found a
small colony of a cleistothecial ascomycete on a diluted
plate of dried powder of salmon and 'wakame' (undaria pin-
natifida (Harvey) Suringar) for cooking of a rice ball. The
fungus occurred on the plate of potato-dextrose agar (PDA)
with 20% sucrose at 25°C and was isolated on the PDA slant
medium. After observation on the media suitable for its
growth, we determined that this fungus is a typical member
Of Coniochaetidium in the Coniochaetaceae (Ascomycetes).
It is described as a new species of the genus.
Coniochaetidium mirabile Udagawa & Tsubouchi, sp.nov.
(Figs. 1-6)
Stat. Anam. Cladobotryum.
Coloniae in agaro farina avenacea restrictae, plus
minusve floccosae, irregulariter sulcatae, tenues, primo
albae, deinde roseo-vinosae vel dilute rubrae; ad centrum
ascomata abundanter producta, cinerascentia; reversum
roseo-bubalinum vel testaceum.
64
Ascomata superficialia, in catervis parvis aggregata,
cinerea, globosa vel subglobosa, 185-280 um diam, saepe ex
hyphis hyalinis, confertis obtecta, nonostiolata. Peridium
ascomatis tenue, ca. 8-10 um crassum, atrobrunneum, bi-
stratum; stratum exterius ex interjectis 'textura intricata'
-'textura angularis' et cellulis incrassatis, brunneis, 3-7
um diam compositum; stratum inferius ex 'textura globulosa'
et cellulis tenuibus, hyalinis, globulosis, 6-20 um diam
compositum. Asci singulares et irregulariter dispositi,
2-8-spori, subglobosi vel ovoidei vel pyriformes, 12-15 x
(8-)10-11 um, tenues, evanescentes. Ascosporae valde oliva-
ceo-brunneae vel atrobrunneae, unicellulares, lenticulares,
5-6(-10) x 4.5-5(-8) x 3.5 um, superficies convexae laeves,
fissura germinali aequatoriali notata et duabus cristis
aequatorialibus ca. 0.5 um latis praeditae.
Mycelium ex hyphis hyalinis, ramosis, septatis, laevi-
bus, 2-4.5 um latis compositum. Conidiophora micronemata
vel semimacronemata, mononemata, suberecta, hyalina, sim-
plicia vel aliquanto subverticillate ramosa, 10-25 x 1.5-
3.5 um, laevia, septata. Cellulae conidiogenae discretae,
singulares vel 2-3(-4) in verticillo disposae, rectae vel
parum curvatae, subulatae, saepe ad medium inflatae et
sursum angustatae, hyalinae, laeves, vulgo 10-28 x 2.5-3 um.
Conidia holoblastica, apice cellularum conidiogeneorum
oriunda, uni-cellularia, hyalina, laevia, tenuia, late
ellipsoidea vel pyriformia vel clavata, 3-7(-10) x 2.5-5
(-6.5) um, superne rotundata et basi anguste truncata, saepe
cicatrice basali instructa, in catenis brevibus fragilibus
cohaerentia.
Holotypus: cultura NHL 2960, ex cibo sicca, Nagoya in
Japonia, in 1984.9 a H. Tsubouchi, isolata. In collectione
fungorum 'National Institute of Hygienic Sciences (NHL),
Tokyo, Japan'.
Etymology: Lat. mirabilis=extraordinary, referring to
the unusual habitat of this fungus.
Colonies on oatmeal agar growing restrictedly, attain-
ing a diameter of 2.7-2.8 cm within 28 days at 23°C, more
or less floccose, irregularly furrowed, composed of a thin
basal felt with scattered patches of dense aerial mycelium,
at first white, then quickly changing to pink color near
Rosy Vinaceous (Rayner, 1970) or pale red (Methuen 8A3 in
Kornerup and Wanscher, 1978); ascomata abundantly produced
at colony center, usually obscured by the overlying aerial
growth, grayish in color; reverse Rosy Buff to Brick
(Rayner) or brownish orange (Methuen 7C4).
Ascomata superficial on the mycelial felt, gregarious
in small clusters, gray, globose to subglobose, 185-280 ym
65
Figure 1. Coniochaetidium mirabile. A. Asci and asco-
spores. B.A portion in cross section of ascoma. C. Coni-
diophores, conidiogenous cells and conidia.
66
in diam, often covered with hyaline, dense hyphal wefts,
nonostiolate. Ascomatal peridium thin, ca.8-10 um thick,
dark brown, two-layered, with a pigmented outer layer
composed of an intermediate between textura intricata and
textura angularis with thick-walled, brown cells 3-7 um in
diam, and inner layer thin, hyaline, of textura globulosa
with globular cells 6-20 um in diam. Asci singly borne on
ascogenous hyphae, irregularly disposed, 2-8-spored, sub-
globose to ovoid or pyriform, 12-15 x (8-)10-11 um, non-
dextrinoid, thin, without apical structures, evanescent.
Ascospores dark olive-brown to dark brown, one-celled,
irregular in shape and size, lenticular, subglobose to
oblate in face view, 5-6(-10) x 4.5-5(-8) x 3.5 um, smooth-
walled, with an equatorial germ slit dividing the spore
into two equal valves, provided with two hyaline equatorial
ridges measuring ca.0.5 um in width, germinating by the
production of one to two germ tubes.
Mycelium composed of hyaline, branched, septate,
smooth-walled, 2-4.5 um diam hyphae. Conidiophores micro-
nematous to semimacronematous, mononematous, suberect,
hyaline, unbranched or branched in somewhat verticillate-
arrangements, 10-25 x 1.5-3.5 um, smooth-walled, septate.
Conidiogenous cells discrete, borne singly or in groups of
2-3(-4), straight or slightly curved, subulate, often
swollen at middle and narrower at the blunt apex, hyaline,
smooth-walled, mostly 10-28 x 2.5-3 um. Conidia holo-
blastic, produced retrogressively as a terminal blow-out at
the tip of individual conidiogenous cells, one-celled,
hyaline, smooth and thin-walled, broadly ellipsoidal to
pyriform or clavate, 3-7(-10) x 2.5-5(-6.5) um, rounded
above and narrowly truncate at the base, often with a prom-
inent basal scar, produced in short fragile chains up to
50 um in length.
Growth at. 37°C is) nil.
Water relations (at 25°C, 14 days): on MY50G, water
activity of 0.89 (Pitt and Hocking, 1985), colonies 5.3 mm
in diam; on MY5-12, water activity of 0.93 (Pitt and
Hocking), colonies 23.1 mm in diam; on MY10-12, water
activity of 0.88 (Pitt and Hocking), colonies 13.8 mm in
diam. Further observation shows in Table 1.
Specimen examined: ex dried salted food (powder of
Salmon and 'wakame' (Undaria pinnatifida (Harvey) Suringar);
the salt contents 38-40%), Nagoya-City, Aichi-Pref., Japan,
September 1984, coll. H. Tsubouchi, NHL 2960 (Holotype) .
Coniochaetidium mirabile can be distinguished from c.
ostreum Malloch & Cain (Malloch and Cain, 1971), the type
species of the genus, by the irregular production of 2 to 8
ERRATA, VOLUME TWENTY-SEVEN
Page 67 line 17 for (%, v/v) read (%, w/v)
67
ascospores per ascus and the anamorph, which is more com-
plexly branched. The ascospores of C. mirabile and Cc. ost-
reum (5.0-8.0 x 4.0-5.5 wm fide Malloch and Cain, 1971)
have a similar range in size, but those of the former have
hyaline equatorial ridges. In this respect of the spore
ornamentation, C. mirabile is very close to two unpublished
collections of Coniochaetidium from dead wood, in the Cryp-
togamic Herbarium of the University of Toronto: TRIC 33447
and TRTIC 45662, both have hyaline equatorial flanges on the
ascospores. However, both the collections are separated
from C. mirabile by their larger ascospore dimensions, cla-
vate and long-stipitate asci, and of a Graphium-like syn-
nematous anamorph.
Table 1. The effect of a on growth of Coniochaetidium
mirabile in PDA medillm containing sodium chloride
NaCl a Colony diam (mm) after incubation at 25°C in
(%, v/v) 1 wk 2 wk 3 wk 4 wk 5 wk
0 0.996 4.2 ia Wo 21.4 (days!
5 oy Co Oe i t\25),0 29.1 31.8 Byan'6|
10 0.938 BG 1 95 28 . 1 32.9 SOnc
15 0.906 15 9.07 14.9 205 25.6
20 Deofemy <0 <0) SROs SS Oca Oa
* First observed time (week) for conidial formation.
** Growth was observed on half of total 12 plates tested.
The Cladobotryum anamorph of C. mirabile is charac-
terized by diffusely verticillate conidiophores and fragile
chains of hyaline, one-celled conidia which are produced
holoblastically from a determinate, retrogressive conidio-
genous cell.
As might be expected from its spoilage of high salt
content food, the fungus grew best on PDA containing 5% and
10% v/v NaCl (see Table 1. However, its salt-tolerance is
not as high for sexual reproduction, showing no sign of any
teleomorph at ay levels below 0.972, despite rather profuse
asexual sporulation within the experimental period.
ACKNOWLEDGMENTS
Appreciation is expressed to Professor D. Malloch of
the University of Toronto for his useful suggestion and
reviewing the manuscript. The authors are also indebted to
Dr. Y. Otani for correcting the Latin diagnosis.
68
Figures 2-6. Coniochaetidium mirabile. 2. Colony on PDA
+ 5% NaCl after 14 days at 25°C. ~<1/2. - 3. A’ portion im
cross section of ascoma. x450. °4. Asci?=x1050.-"5. S-E om:
of ascospores. x1900. 6. Conidiophores, conidiogenous
cells and conidia. x880.
69
LITERATURE CITED
Kornerup, A., and J.H. Wanscher. 1978. Methuen Handbook of
Colour. 3rd ed. Eyre Methuen, London.
Malloch, D., and R.F. Cain. 1971. New cleistothecial
Sordariaceae and a new family, Coniochaetaceae. Can.
J. Bot. 49: 869-880.
Pitt, J.I., and A.D. Hocking. 1985. New species of fungi
from Indonesian dried fish. Mycotaxon 22: 197-208.
Rayner, R.W. 1970. A Mycological Colour Chart. Common-
wealth Mycological Institute, Kew & the British
Mycological Society.
MYCOTAXON
Veue XV EL Dp. knoe October-December 1986
MATERIALS FOR A LICHEN FLORA OF THE ANDAMAN ISLANDS-I
M.B. NAGARKAR, P.K. SETHY & P.G. PATWARDHAN
Department of Mycology, M.A.C.S. Research Institute, Pune-411 004, India.
ABSTRACT
Eighteen species of Thelotremataceae from the Andaman Islands, India
are described. Leptotzema pertusarioides, Ocellulartia wandoorensts and
Thelotrema guptei are new species. L. nuwarense, O. olivacea, O. xanthost-
10miza and T. 12cutvum are additions to the lichen flora of India.
INTRODUCTION
Oceanic islands have always attracted the attention of floristic botanists.
In his famous lecture on "Insular Floras", delivered at Nottingham on 27th
August 1866, Joseph Dalton Hooker remarked, "The relationships between these
oceanic island Floras are of two kinds.....One a relationship of analogy between
themselves, due to physical conditions common to them all- to their climate,
exposure, limited area, distance from continents etc. Thus they are rich in
Ferns, Mosses and other Flowerless plants.....Species are few in proportion to
genera, and genera in proportion to the orders.....The other is a relationship of
affinity, a bona fide kinship, which the Floras of islands display in common
with one another or with certain continents." (Williamson, M. 1984).
The Andaman islands attracted the attention of floristic botanists as
early as 1791 (Thothathri 1960) but lichens were first collected from these
islands in 1867 by Dr.Kurz of the Royal Botanic Garden, Calcutta.
Reports of 62 taxa identified by Nylander (1873), 24 taxa by Jatta (1905)
and 47 species of foliicolous lichens by Ajay Singh (1979) form the major cont-
ribution to our knowledge of the lichens of the Andaman islands. Ajay Singh
(1979) has listed all the taxa reported so far and reviewed earlier work on
lichens from these islands.
In 1985 we made lichen collections on the Andaman islands under a
project entitled,"Lichen flora of the Andaman islands" sponsored by the
Department of Science and Technology, New Delhi, India. We propose
TZ
to describe and revise the lichen taxa from these islands in a series of
papers, as we complete studies of the various families.
The Andamans constitute a group of islands lying in the Bay of Bengal
between 13° 41' and 10° 30' N latitude and 92° 11' and 93° 7' E longitude,
1100 km from the Coromandel! coast of the Indian mainland and 515 km from
the Tenasserim coast of Burma. The land area of the entire group of 204
islands is about 6330 km”. The main group consists of North Andaman, Middle
Andaman and South Andaman. Little Andaman is located south of the main
group (Great) of the Andamans.
The vegetation of the islands is classified under six types (Thothathri
1962):
. Mangrove forest (confined to creeks)
. Beach forest (on the coast and extending somewhat inland)
- Deciduous forest
. Evergreen forest at lower elevations
- Moist evergreen forest at higher elevations
. Vegetation in cleared areas
NUOFWNe
In his studies on the flora of the Andaman islands Thothathri (1960)
concluded, "In general it may be summed up that the flora of the Andaman
Islands is an admixture of Burmese and Malayan plants together with its
indigenous plants.'"(p. 361).
In this paper we deal with 18 species of the four ascospore-based
genera Leptotiema, Ocellulatia, Phaeottema and Thelottema in the lichen
family Thelotremataceae. Several others remain to be studied. Distribu-
tion records outside of the islands are taken from Hale (1974, 1978, 1981),
from our own identified materials in AMH and from examination of types.
Leptotrema elachistoteron (Leighton) Patw.& Kulk., Now. J. Bot., 24:
128 (1977). (Fig. 1)
Thelotrema elachistoteron Leighton, Ttans. Linn. Soc. London, 27: 169(1869).
Thallus pale yellowish gray to straw coloured, smooth, uneven, medulla
50-100 tum thick, epiphloeodal; apothecia many, solitary, immersed to slight-
ly raised, 0.2-0.4 mm diam; ostiole minute, round, slightly raised dark brown
rim. 80-130 um across; exciple fused, colourless to reddish brown at the apex
only; hymenium 100-150 um high, 200-250 um broad; ascospores brown, muri-
form, 2-4 per ascus, 15-20 X (42) 60-85 pm, I + violet.
Chemistry: Thallus K + deep yellow, P + orange; stictic and constictic
acids present.
Specimens examined: South Andaman, Baratang island, Lorojig - 85.237,
262, 264, 227.
Distribution: Sri Lanka, India.
Leptotrema zeclusum (Krempelh.) Zahlbr., a comparable species, has
larger ascospores exceeding 100 ym in length.
(ES:
74
Leptotrema nuwarense (Hale) comb. nov. (Fig.#2)
Myriotrema nuwartense Hale, Bull. Bt. Mus. nat. Hist. (Bot.) 8 (3): 289 (1981).
Thallus pale greenish gray, uneven, cracked, medulla 100-130 pm thick,
crystalloid, partly hypophloeodal; apothecia numerous, solitary, immersed,
0.2-0.35 mm diam; ostiole minute, round, off-white annulate rim, 180-250 um
across; exciple free, inner exciple forming a ring, colourless; hymenium
100-130 pm high, 200-250 pm broad; ascospores brown, muriform, 2 per ascus,
10-15 X 42-55 ym, I-.
Chemistry: Thallus K + yellow, P + orange; stictic and constictic acids
present.
Specimens examined: South Andaman, Wandoor, Alexandria island -
85.1108b, 1035, 1104, 1106.
Distribution: Sri Lanka, India
Leptotrtema pertusarioides, sp. nov. (Fig. 3)
Thatlus corticola, pallide fulvo-cinereus, vertucosus, Crassus, pro parte
hypophfoeodes; apothecia numerosa, congesta, emergentia, basin constucta,
0.6 - 0.85 mm diametro; ostiolum totundatum, apertum, depressum, 200-400
um diametro; excipulum connatum, non-cartbonaceum; columella nulla; hyme-
nium 200-250 wm altum; sporae octonae, fuscae, late elliosoideae, 4-locula-
tae, 10-12 X 15-18 um.
Thallus pale yellowish gray, warty, medulla 160-250 ym thick, crysta-
lloid, partly hypophloeodal; apothecia numerous, crowded, emergent, const-
ricted at the base, 0.6-0.85 mm diam; ostiole open, depressed, 200-400 pum
across; exciple fused, reddish yellow; hymenium 200-250 wm high, 250-320
pm broad; ascospores brown, muriform, 8 per ascus, broadly ellipsoidal with
1 transverse septum in the middle and | faint vertical septum forming 2
locules in each half, 10-12 X 15-18 pm.
Chemistry: Thallus K + yellow, P + blood red; protocetraric acid
present.
Typus: India, Andaman islands, South Andaman, Wandoor, Alexandria
island, leg. M.B. NAGARKAR and P.K. SETHY, 6 March 1985, 85.1115
(AMH-holotypus; US-tsotypus).
This species has apothecia extremely similar to those of the genus
Pertusatia, but the spores are typically broadly ellipsoid with one distinct
transverse septum and two locules in each compartment.
Leptotrema bahianum (Ach.) Mull. Arg., another species with proto-
cetraric acid, has somewhat larger ascospores (12-14 X 16-24 um) with
5-6 transverse and 2-3 longitudinal locules.
Leptotiema wecdusum (Krempelh.) Zahlbr., Cat. Lich. Untv., 2: 639 (1924).
(Fig. 4)
Thelotrema 1eclusum Krempelh. in Nyf., Bull. Soc. Linn. Nowmand., 7(2): 168
(1873).
Thallus brownish gray, rough, becoming cracked in mature stage,
medulla 150-200 pm thick, crystalloid, epiphloeodal; apothecia numerous,
solitary, immersed to slightly raised, 0.3-0.5 mm diam; ostiole round, mode-
is
rately open, 100-200 um across; exciple fused, pale yellow; hymenium 180-220
ym high, 250-310 pm broad; ascospores brown, muriform, 1 per ascus, 18-32 X
(73)96-126 pum, I-.
Chemistry: Thallus K + yellow, P + orange, stictic and constictic acids
present.
Specimens examined: South Andaman, Wimberliganj, Kalatang- 85.1217,
Wandoor, Alexandria island- 85.1111.
Distribution: Panama, India.
Ocellularia allosporoides (Nyl.) Patw.& Kulk., Kavaka, 5: 5 (1977). — (Fig. 5)
Thelotrema allosporoides Nyl., Bull. Soc. Linn. Notmand., 7(2): 167 (1873).
Thallus greenish gray to yellowish gray, smooth to rough, medulla 160-
200 um thick, corticated, epiphloeodal; apothecia numerous, solitary, semi-
emergent to emergent, 1.2-1.8 mm diam; ostiole depressed, open, 360-630 jum
across; exciple carbonized laterally; columella simple, tip white pruinose,
carbonized, 400-600 pm thick; hymenium 200-250 jm high; ascospores hy al-
ine, transversely septate, 4 per ascus, 12-15 X 96-128 um, I + deep violet.
Chemistry: Thallus K-, P-; lacking lichen substances.
Specimens examined: Andaman islands, South Andaman, Chidya T apu-
85.52, 59, 61, 69, 74(b), 75. Wimberlig anj, Kalatang- 85.82, 89, 98, 105, 110,
PS 1927 19355 154,200, 1215, 1216; 'Alexandriiaisiand-'$5.1038,5 (1093, 1127,
1142.
Distribution: India, New Zealand.
Ocellulatia berkeleyana (Mont.) Zahlbr., Englet-Prantl, Naturl. Phlanzenfamil.,
1. Teil, Abt: 137 (1905). (Fig. 6)
Stegobolous berkeleyanus Mont. in Hook. London J. Bot., 4: 4 (1845).
Thallus light greenish gray, smooth, medulla 20-40 tum thick, corticated,
epiphloeodal; apothecia many, solitary, emergent, constricted at the base,
0.6-1 mm diam; disc wide open, white pruinose, 300-600 ym across; thalline
exciple erect, proper exciple carbonized laterally; columella actinoid with
3 to 4 branches, prominent from the exterior, protruding out as a white pru-
inose tip, carbonized; hymenium 70-90 ym high; ascospores hyaline, transvers-
ely septate, 8 per ascus, 3-5 X 12-15 um, 4 loculate, I + violet.
Chemistry: Thallus K -, P + Yellow; psoromic acid present.
Specimen examined: South Andaman, Wandoor, Alexandria island- 85.1065.
Distribution: Guadeloupe, Panama, Cuba, Apiahy, Brazil, India, Java,
Philippines, South Africa
Ocellulatia diacida Hale, Mycotaxon, 7(2): 378 (1978). (Fig. 7)
Thallus olivaceous green to greenish gray, smooth, medulla 20-35 um
thick, corticated, hypophloeodal; apothecia numerous, solitary, semi-emergent,
0.5-0.75 mm diam; ostiole open, often filled with tip of the columella, brown
to dark coloured rim, 147-210 um across; exciple carbonized in the ostiolar
region only, sometimes extend further; columella simple, distinct, formed of
thick walled hyphae apically, 120-190 yum thick, carbonized; hymenium 80-110
lum high; ascospores hyaline, transversely septate, 8 per ascus, 4-7 X 16-32 jumy
76
(3)5 - &(10) loculate, I + violet.
Chemistry: Thallus K + pale yellow, P + pale yellow; 3-4 spots above
norstictic level in TLC, 'diacida' unknown.
Specimens examined: South Andaman, Port Mount - 85.33, Wandoor, Alex-
andria island - 85.1056, 1059, 1074, 1075, 1094, 1097, 1108,1136.
Distribution: India, Sri Lanka
Ocellularia olivacea (Fée) Mill. Arg. in Mém. Soc. Phys. Hist. nat. Geneve,
29(8): 7 (1887). (Fig. 8)
Myttotrema olivaceum Fée, Essai Crypt.: 103 (1824).
Thallus light greenish gray to ashy gray, smooth, deeply fissured, areolate,
medulla 250-380 um thick, epiphloeodal; apothecia numerous, solitary, immersed,
0.15-0.25 mm diam; ostiole round, slightly raised off-white annulate rim, 50-100
yum across; exciple free, thin, pale reddish, inner exciple forming a minute pore
within the main pore; hymenium 70-90 pm high, 140-160 um broad; ascospores
hyaline, transversely septate, 8 per ascus, 5 X 12.5-15 ym, 4 loculate, I + violet.
Chemistry: Thallus K -, P -3 'olivacea' unknown present.
Specimen examined: South Andaman, Baratang island, Lorojig - 85.222.
Distribution: West Indies, Central and South America, Sri Lanka, New
Caledonia, Marquesas islands.
The present species is comparable in ascocarp morphology and anatomy
to Ocellulatia mictopora (Mont.) Mull. Arg. which contains psoromic acid and
Ocellularia glaucula (Nyl.) Zahlbr. which contains hypoprotocetraric acid.
Ocellulatia pycnophtagmia (Nyl.) Zahlbr., Cat. Lich. Univ. 2: 599 (1924).
(Fig. 9)
Thelotrema pycnophtagmium Nyl., Settum Lichen. Tropic Labuan et Singapore.
Pe Datta).
Thallus whitish gray, smooth, continuous, medulla 50-80 jm thick, crys-
tals, mostly hypophloeodal; apothecia many, scattered, solitary, rounded to
somewhat elongated, semiemergent, chroodiscoid, 0.6-1.2 mm diam; disc wide
open, white pruinose; exciple free with periphysoids, thalline rim recurved,
non-carbonized; hymenium 150-200 um high, 500-800 pm broad; ascospores
hyaline, transversely septate, 4-8 per ascus, 5 - 7.5 X 70 - 97.5 yum, 25-35
loculate, I + deep violet.
Chemistry: Thallus K-, P-; lacking lichen substances.
Specimens examined: South Andaman, Baratang island, Nilambur- 85.489,
Wrafter's Crick- 85.568.
Distribution: Labuan, India
Ocellularia terebrans (Nyl.) Zahlbr., Cat. Lich. Univ. 2: 602 (1924). (Fig. 10)
Thelotrema terebtans Nyl., Bull. Soc. Linn. Nowmand., 7(2): 166 (1873).
Thallus yellowish gray to straw coloured, shiny, warty, medulla 360-450
yum thick, studded with crystals, epiphloeodal; apothecia numerous, solitary,
immersed, 0.25 - 0.35 mm diam; ostiole open, flush with the thallus, round,
white rimmed, 150-200 yum across; exciple free, pale yellowish, inner exciple
separated forming an inner ring, periphysoids; hymenium 120-150 pm high,
- .
NG a
|
78
200-240 um broad; ascospores hyaline, transversely septate, 8 per ascus, 4.8 X
19-25(30) pm, 7-10 loculate, I + pale blue.
Chemistry: Thallus K + deep yellow, P + orange; stictic acid present.
Specimen examined: South Andaman, Chidya Tapu- 85.63.
Distribution: India, Sri Lanka.
Ocellulatia wandoorensis, 4). nov. (Fig. 11)
Thatlus corticola, viridi-cinerteus, veriucosus, hypophloeodes; apothecia
numerosa, solitaria, arcte disposita semi-emergentia vel emergentia, 0.5-0.8
mm diametro; ostiolum totundatum apertum, 0.2-0.3 mm diametto; exctoulum
connatum, intus catbonaceum; columella actinoidea, 3-4 furcata, badia vel
fusca, apice epruinosa vel leviter albo-pruinosa; hymentum 50-70 pm altum;
sporae octonae, incolores, transversm septatae, 4 loculatae, 3-5 X 8-10 pm,
1 + coeruleae.
Thallus greenish gray, warty, medulla 60-30 um thick, corticated, hypo-
phloeodal; apothecia numerous, solitary, closely disposed, semi-emergent to
emergent, 0.5-0.8 mm diam; ostiole rounded, open, pore gaping, 230-300 pum
across; exciple carbonized laterally; columella actionoid with 3 to 4 branches,
reddish brown to dark brown, tip epruinose to lightly white pruinose; hymenium
50-70 um high; ascospores hyaline, transversely septate, 8 per ascus, 3-5 X
8-10 pum, 4 loculate, I + violet.
Chemistry: Thallus K-, P-; lacking lichen substances.
Typus: India, Andaman Islands, South Andaman, Wandoor, Alexandria
island, leg. P.K. Sethy and M.B. Nagarkar, 6 March 1985, 85.1103 (AMH-
holotypus, US- tsotypus).
Other specimens examined: South Andaman, Alexandria island, wandoor-
85.1080, 1102.
This species resembles to Ocellularia betkeleyana (Mont.) Zahlbr. in
ascospore morphology. O. berkeleyana however, has a very prominent protru-
ding columella with a heavily pruinose tip. The ostiolar region of the exciple
and the disc are also pruinose, unlike O. wandoorensis. O. glaziovu Mill. Arg.
has an even more highly developed actionoid disc.
Ocellulatia xanthostromiza (Nyl.) Zahlbr., Cat. Lich. Univ. 2: 604 (1924).
(Fig. 12)
Ascidium xanthostromiza Nyl., Settum Lichen. Tropic. Labuan et Singapore:
20 (1891).
Thallus greenish mineral gray, smooth to rugulose, cracked, medulla
150-180 wm thick, large crystals, partly hypophloeodal; apothecia numerous,
crowded, emergent, constricted at the base (ascidioid), 0.6-0.8 mm diam;
ostiole carbonized laterally; columella simple, 150-250 yum thick, carbonized;
hymenium 110-130 pum high; ascospores hyaline, transversely septate, 8 per
ascus, 3-5 X 11-20 pm, 3-6 loculate, I + reddish violet.
Chemistry: Thallus K + yellow, P + red; 'cinchonarum' unknown present;
medulla pale orange.
Specimen examined: South Andaman, Wimberliganj, K alatang-85.182.
Wandoor, Alexandria island- 85.1147.
Distribution: Singapore, India.
79
This species is close to Ocellulatuia cavata (Ach.) Miill. Arg. in the peculi-
ar chemistry and morphology of ascocarps but O. cavata has larger ascospores
(6-8 X 25-43 pm).
Phaeotzema facteum (Krempelh.) Mill. Arg. in Flota, Jena, 70: 398 (1887).
(rigs 3)
Thelotrema facteum Krempelh. in Nyl. in Floza, Jena, 47: 269 (1864).
Thallus whitish to pale yellowish, smooth, medulla 15-30 ym thick, hypo-
phloeodal; apothecia numerous, solitary, emergent, 0.6-0.85 mm diam; disc
wide open, white to ashy white pruinose, 200-500 pm across; thalline exciple
suberect to erect, inner exciple pulling away from the thalline exciple, free,
reddish brown; hymenium 140-160 pm high, 300-550 pm broad; ascospores
brown, transversely septate, 4 per ascus, 10-12 X 37-50 pm, 10-15 loculate.
Chemistry: Thallus K-, P-3 lacking lichen substances.
Specimen examined: South Andaman, Baratang island, Nilambur-85.484.
Distribution: United States, West Indies, Central America, Kenya,
Mozambique, South Africa, Sri Lanka, Japan, New Guinea, Australia, Solomon
islands, New Caledonia, Polynesia, Hawali islands, India.
Thelotrema glaucopallens Nyl., Ann. Sci. nat. (Bot.) IV, Il: 327 (1863).
(Fig. 14)
Thallus light greenish gray, smooth, shiny, medulla 330-400 pm thick,
studded with crystals, corticated, epiphloeodal; apothecia numerous, solitary,
immersed to slightly raised, 0.2-0.35 mm diam; ostiole round, flush to barely
raised, off-white to pale coloured annulate rim, 120-170 pm across; exciple
fused, non-carbonized to weakly carbonized at the apex only; hymenium 90-
110 pm high, 150-180 pm broad; ascospores hyaline, muriform, 8 per ascus,
7-9 X 22-25 (29.4) pm, 1-3 X 7-10 loculate, I-.
Chemistry: Thallus K + yellow, P + orange; stictic and constictic acids
present.
Specimens examined: South Andaman, Chidya Tapu- 84.51, 66, 74(a),
Wimberliganj, Kalatang- 85.94, 128.
Distribution: Panama, Dominica, Guatemala, Guyana, Venezuela,
Trinidad, Tanzania, Ivory Coast, Union of South Africa, Philippines, India.
Thelotzema guptei, sp. nov. (Pigs 15)
Thallus albo-cinereus, inaequalis, crassus, pro parte hypophtoeodes;
apothecia tara, dispersa, solitaria, semi-emergentia vel emergentia, 0.8 - 1.3
mm diametto; ostiolum rotundatum, apertum, 0.35-0.45 mm diametro; exctpu-
lum connatum, intus catbonaceum; columella cylindrica, carbonacea, 80-120
pm crassa; hymenium 350-400 pm altum; sporae 1: nae, incolores, murales,
21-33 X 121-172 pm, 1 + caewlescentes.
Thallus whitish gray, uneven, cracked, medulla 60-90 pm thick, crys-
talloid, corticated, partly hypophloeodal; apothecia few, scattered, solitary,
semi-emergent to emergent, 0.8-1.3 mm diam; ostiole open, 350-450 ym across;
exciple carbonized; columella simple, 80-120 pum thick, carbonized; hyme-
80
350-400 ym high; ascospores hyaline, muriform, | per ascus, 21-33 X 121-172
ym, I + deep violet.
Chemistry: Thallus K-, P-; lacking lichen substances.
Typus: India, Andaman islands, South Andaman, Port Mount, leg. P.G.
Patwardhan and M.B.Nagarkar, 14 February 1985, 85.21 (AMH- holotypus,
US- tsotypus).
Specimen examined: South Andaman, Port Mount- 85.21.
Thelotrema depressum Mont., the only comparable species has larger
( upto 2 mm diam. ), heavily carbonized ascocarps constricted at the base.
This new species is named after Mr. S.C.Gupte, former Conservator of
Forests, Andaman and Nicobar circle.
Thelotrema massatongot (Mont.) Zahlbr., Cat. Lich. Untv., 2: 624 (1924).
(Fig. 16)
Ascidium massatongot Mont., Ann. Set. nat. (Bot.) IV, 14: 174 (1860).
Thallus pale yellowish gray, smooth to finely warty, medulla 100-120 jum
thick, studded with crystals, dense cortex, epiphloeodal; apothecia numerous,
solitary to group of 2 to 4, strongly emergent, constricted at the base, asci-
dioid, 1-1.5 mm diam; ostiole round, depressed, surrounded by a strongly raised
rim, 400-700 um across; exciple carbonized laterally; columella simple, base
broader than the apex, tip white to off-white pruinose evident through the
pore, 120-200 pm thick; hymenium 300-400 pm high; ascospores hyaline, muri-
form, 1 per ascus, 25-33 X 127-207 pum, I + deep violet.
Chemistry: Thallus K + yellow, P + orange; salacinic acid present.
Specimens examined: South Andaman, Baratang island, Wrafter's Crick-
85.646, 650, 651, 652, 653.
Distribution: India, Sabah, Sri Lanka, Malaya, Philippines, Japan, Solomon
islands, Australia
Thelotrema poltillense Vainio in Suom.Tiedeakat. Toim. A, 15(6): 180 (1921).
(Fig. 17)
Thallus pale greenish gray, smooth medulla 60-100 pum thick, epiphloeo-
dal; apothecia numerous, solitary, semi-emergent to emergent, 0.8-1.2 mm
diam; ostiole open, brown rim, pore gaping, 350-500 jm across; exciple car-
bonized laterally; columella very wide, 400-600 pm thick, carbonized, becoming
divided at maturity, no hymenium part develop in the dividing region, white
pruinose at the tip; hymenium 70-90 pm high; ascospores hyaline, muriform, 8
per ascus, 4.8-6.4 X 11-17 pam, 4-6 transverse locules, 2 longitudinal locules,
I + violet.
Chemistry: Thallus K-, P + yellow; psoromic acid present.
Specimen examined: South Andaman, Wimberliganj, K alatang- 85.79.
Distribution: Sri Lanka, New Caledonia, Philippines, India.
The columella in the Andaman material is vertically furrowed but no
asci or paraphyses develop in the furrows.
Thelotrema recutvum G. Salisb., Rev. Bryol. et Lichenol., 38: 285 (1972).
(Fig. 18)
81
Thallus whitish gray, smooth, medulla 30-50 um thick, crystalloid, partly
hypophloeodal; apothecia numerous, solitary, rounded to elongated, semi-
emergent, chroodiscoid, 1-2.5 mm diam; disc wide open, white to ashy pruinose;
exciple free, periphysoids, thalline margin recurved, non-carbonized; hymenium
100-140 um high, 600-1200 jum broad; ascospores hyaline, muriform 2/4 per
ascus, 15-20 X 40-68 pm, I + deep yellow.
Chemistry: Thallus K-, P-; lacking lichen substances.
Specimen examined: Andaman island, South Andaman, Wandoor, Alexa-
ndria island- 85.1146.
Distribution: Philippines, India.
ACKNOWLEDGEMENTS
We are grateful to Mr. S.C.Gupte, Ex. Conservator of Forests, Andaman
and Nicobar Islands, India for his active interest in the field work and to
Dr. Mason E.Hale, Smithsonian Institution, Washington D.C. for critically
going through the manuscript and valuable suggestions.
We are also grateful to Department of Science and Technology, New
Delhi for financial support.
LITERATURE CHED
Hale, M.E. 1974. Morden-Smithsonian Expedition to Dominica: The Lichens
(Thelotremataceae). Smithsonian Contr. Bot., 16: 1-46.
Hale, M.E. 1978. A Revision of the Lichen Family Thelotremataceae in
Panama. Smithsonian Contr. Bot., 38: 1-60.
Hale, M.E. 1981. A Revision of the Lichen Family Thelotremataceae in
Sri Lanka. Bull. Bt. Mus. nat. Hist. (Bot.), 8 (3): 227-332.
Jatta, A. 1905. Licheni Esotici dell'Erbario Lavier reccolti nell'Asia
meridionale, nell'Oceania, nell'Biasile e nell Madagascar. Malpighia,
19: 162-186.
Nylander, W. 1873. Lichenes insularum Andaman. Bull. Soc. Linn.Notmand.,
Hy 72 '62-183.
Singh, A. 1979. The lichen flora of India with special reference to Andaman
Islands. Pt0g1ess in Plant Research, Silver Jubilee Publication, N.B.R.1.,
1: 39-56 (T.N. Khoshoo and P.K. Nair, Eds.).
Thothathri, K. 1960. Studies on the flora of the Andaman Islands. Bull.
bot. Surv. India, 2 (3 & 4): 357-373.
Thothathri, K. 1962. Contributions to the flora of the Andaman and Nicobar
Islands. Bull. bot. Surv. India, 4 (1-4): 281-296.
Williamson, M. 1984. Sir Joseph Hooker's Lecture on Insular Floras. Biol. J.
Linn. Soc., 22: 59-77.
SZ
Explanation of figures 1-18: 1. Leptotzema elachistoteton (85. 237); 2. L.
nuwatense (85. 1108b); 3. L. pertusarioides (85. 1115, Holotype-AMH); 4.
L. reclusum (85. 1217); 5. Ocellularia allosporoides (85. 59); 6. O. berketeyana
(85. 1065); 7. O. diacida (85. 33); 8. O. olivacea (85. 222); 9. O. pycnophrtagmia
(85. 489); 10. O. terebrans (85. 63); 11. O. wandoorensts (85. 1180, Holotype-
AMH); 12. O. xanthostromiza (85. 182); 13. Phaeotrema facteum (85. 325);
14. Thelotrema glaucopallens (85. 51); 15. T. gupteé (85.21, Holotype-AMH);
16. T. massalongos (85.646); 17. T. politlense (85. 79); 18. T. tecutvum (85.
1146). Scale in mm.
MYCOTAXON
Vol. XXVIE, pp..963-86 October-December 1986
Hyphomucor, a new genus in the Mucorales for
Mucor assamensis.
Mths Ar Co) e rs
Centraalbureau voor Schimmelcultures,
Ve Ole bOxX ‘ja, 3740 (RG eBaarn.sttne Netherlands :
ihnree strains tdentified as Mucor assamens is
bDygies Joyce Lunn of “CMD were "studred. “Unlike
MuGerr spp, these "isolates produced a Llaverivof
aerial hyphae on which sporangiophores were
formed; the latter heavely branched with
almost perpendicular branches.
The production of sporangiophores from aerial
hyphae was not mentioned in the original
description. The strains are otherwise
Sinioar (vor'Mucorm- assamensis. tire stoype
cubvure "of Mucor -assamensis, 1so0lated from
forsst (soll on Tezpur, ASsam’, Was. not
available.
A new genus is proposed to accomodate "Mucor
assamensis".:
Hyphomucor Schipper & Lunn, gen. nov.
Sporangia multispora, anapophysata,
columellata: sporangiopnora ‘ramosa, -oriuntur
ex~phyphae ‘aeriae vel superficialibus,
zygosporae ignotae.
type species Hyphomucor assamensis (Mehrotra
& Mehrotra) (Schipper & bunk =! Mucor
assamenous *B.o. Mehrotra GB. Ri. Menrotra: —
Sydowia 23: 184. 1969.
Multispored, non-apophysate, columellate
sporangia are borne on branched
sporangiophores, arising from aerial hyphae
and superficial hyphae. Zygospores are
unknown.
84
The genus differs from Mucor by ete
produce tion "of “sporangirophores” or aciria
hyphae, from Mortierella Dy drstinciuly
columellate sporangia,, from Thermomucor and
Rhizomucor in the absence of growth above
36,02 The cenus 2s distineuresved mi rom
AbSidia, Ac tinomucor -andihiaZopus 4by- vie
absence “OVC STING E ESO LOnsS Vander Zolds.
Simple sporangiophores as seen in Rhizopus
and apophysate (sporangia vas. seen tm AbSidia,
dO MOCHOCCUr win AY phoOmMmico Tr .
Hyphomucor assamensis, (Mehrotra & Mehrotra)
SGHUD Ger “tun COMD.. Ow,
= Mucor ,assamensis B.S. Mehrotra .¢ \.B.R. Mehrotra:
The, following description. 1s ibased: on sCBs
WAS P71 6 On, Uihwtea tse hms
Colony gcreyaush,. With abundant \white, sterile,
aerial mycelium and bDlackish sporangia.
SPpOrangiophores: uitraedent, ore cecurved.. arnais mae
from thin, branched, aerial hyphae, rarely
from superticral hyphae, @lassy white to
brownish, darkest just below the ‘Sporangiua,
repeatedly branched sympodially, narrow at
the base, widening towards the sporangium,, up
to 20 um wide. Sporangia globose, mostly .up
tor fo: UnNedaam,) Occast ona liye vow toy (0 0r e
white to. biack, either dryrand powdery 2n
appearance dorm surrounded ‘by mous ture.
Columelblaetsessile,) applanate or condical,
some: with a‘ bbunt projection sath the cop,
bluish. cod tars .sma ll. .orm-absent).
Sporangiospores sub) cloboséie silitenil y
roughened in outline, with granular convents,
4-6 (8)um diam.
Zygospores unknown.
Influence of temperature: no growth at 36,06
and, above, nor at 6.Gvand below. .Growen,
nO; SpOrular1on 2ti 33) awe rowtey and
sporulation between 12 and Sie optimal
at About 24” CO.
Influence of medium: mature sporangia were
produced. on, beerwort, agar, PDA, cherry
agar, V8-agar and :OA.,.Colonies on PDA at
24°C were, rather .difruse,..alvowing
observation of sporangiophore development.
Pe,
ps
Hyphomucor assamensis comb.
as sporangpiosphores.—
GUO G SS
85
D. columellac.= c. sporangrospores:
Influenceiof Light: \insipniticanv.
Material examined
OBS): 4-15 o fie Se lM 270192 ane ena Ss olhave
identification (G. Satyanarayana)
CBs 54. Co." Mie 13888) “ex yBurmann2aecpy,
Kuala Lumpur, )Malay suai.
Ariffan «supose,
OBSi2535:65) © VME 32 Oi, CxS Oa leis tort (ie saree
(Drs. Coomaraswamy, Subi5)
A fourth strain, Ls0baced, from tropicat
forest soil, Srilanka, by RN. de smonsecaywas
available as an herbarium specimen only (IMI
ZOU Oy MUS OR) Vax Siew)
Mating experiments
No sexual reactions were observed in
contrasts between the strains on various
media and temperatures.
Discussion
Hyphomucor assamensis resembles Thermomucor
indicae-seudaticae, but differs in being
mesophilic’ and in the production of non-=
apophysate sporangia on sporangiophores
arising from aerial hyphae.
Acknowledgements
I) Want .toUtnankeDr Ae fOn. Ons samira) eager ie
Kirk, CML, Rormimeading thetimanuscrip hans.
JeAy Stalpersy tor iprepariwng tie arin
diagnosis.
MYCOTAXON
Vol. XXVII, pp. 87-92 October-December 1986
METARHIZIUM FLAVOVIRIDE VAR. MINUS,
VAR. NOV., A PATHOGEN OF PLANT- AND
LEAFHOPPERS ON RICE IN THE PHILIPPINES
AND SOLOMON ISLANDS
Michiel C. Rombach*
Insect Pathology Resource Center
Boyce Thompson Institute at Cornell University
Tower Road, Ithaca, New York 14853, USA
Richard A. Humber
USDA-ARS Plant Protection Research Unit
Boyce Thompson Institute at Cornell University
Tower Road, Ithaca, New York 14853, USA
Donald W. Roberts
Insect Pathology Resource Center
Boyce Thompson Institute at Cornell University
Tower Road, Ithaca, New York 14853, USA
Metarhizium flavoviride Gams & Rozsypal var. minus var. nov. is described
from the brown planthopper, Nilaparvata lugens Stal (Homoptera, Delphacidae),
on rice in the Philippines and the Solomon Islands and from the zig-zag
leafhopper, Recilia dorsalis (Motschulsky) (Homoptera, Cicadellidae), in the
Philippines. The new variety has also been collected on a grasshopper in the
Galapagos Islands.
The conidia of the new variety are smaller (mostly 4.5-7 x 2-3 um) and more
consistently ellipsoidal to ovoidal than those of M. flavoviride var. flavoviride
[(6.5-)7-9[-11] x 4.5-5.5 [um]. The new variety may form synnemata in culture.
At present, three species of Metarhizium are recognized: Most authors
have followed Tulloch (1976) in accepting M. anisopliae (Metschn.)
Sorok. (with cylindrical phialides bearing chains of cylindrical conidia that
usually adhere laterally to form columnar conidial columns) and M. flavo -
viride Gams & Rozsypal (with narrowly clavate phialides bearing chains
* Seconded to Department of Entomology, International Rice Research Institute, P.O.
Box 933, Manila, Philippines.
88
of ovoid or ellipsoidal conidia that typically adhere in parallel masses).
Rombach et al. (1986b) are restoring the name Metarhizium album Petch
for a pathogen of Asiatic homopterans that forms clavate phialides with
nonadherent chains of white to brown, ellipsoid to ovoid conidia.
Metarhizium flavoviride is primarily an entomogenous species reported
from curculionid beetles (Gams & Rozsypal 1973; Marchal 1977), from
a grasshopper (Evans & Samson 1982), and from agricultural soils in
Germany and the Netherlands (Gams & Rozsypal 1973). This species
has been studied as a specific pathogen of black vine weevils, Otior -
rhynchus sulcatus F. (Marchal 1977; Soares et al. 1983; Poprawski et al.
1985a) and of the onion root maggot, Delia antiqua (Meigen) (Poprawski
et al. 1985b). The small-spored variety described below causes natural
epizootics of the brown planthopper, Nilaparvata lugens Stal, and of the
zig-zag leafhopper, Recilia dorsalis (Motschulsky), on rice in the Philip -
pines and Solomon Islands. Field populations of brown planthoppers can
be infected by sprayed applications of the small-spored variety (Rombach
et al. 1986a).
Metarhizium flavoviride Gams & Rozsypal (1973) is characterized by
the formation of sporodochia bearing clavate phialides (9-14 x 3-4.5 um)
and chains of large ellipsoidal conidia (Fig. 1a); the conidial masses of
this species are pale green to distinctly yellow-green. Cultures of
Metarhizium flavoviride are slow growing, flat, and slow to produce
conidia. The ovoid to ellipsoid conidia and clavate phialides are the major
diagnostic characters which separate M. flavoviride from M. anisopliae
with its distinctly cylindrical conidia and cylindrical phialides.
In recent years, numerous isolations identified as Metarhizium flavo -
viride were made from plant- and leafhoppers on rice in Asia (Fig. 2a).
Most of these isolates are preserved in liquid nitrogen at the Collection of
Entomopathogenic Fungi (ARSEF), USDA-ARS Plant Protection
Research Unit (Boyce Thompson Institute, Ithaca, New York).
On the plant- and leafhoppers (Fig. 1b), Metarhizium flavoviride
shows a slightly different morphology from that originally described for
M. flavoviride by Gams & Rozsypal (1973), most notably in the smaller
dimensions (Table 1) and more consistently ovoid shape of the conidia.
In artificial culture (Fig. 1c), the characteristics are slightly more variable.
Tulloch (1976) separated the varieties M. anisopliae var. anisopliae and
M. anisopliae var. majus (Johnston) Tulloch* mainly by the shape and
dimensions of their conidia. It is appropriate, therefore, to expand the
concept of M. flavoviride defined by Gams & Rozsypal (1973) to accom -
modate these Asiatic isolates from plant- and leafhoppers on rice in a new
* This is an obligatory orthographic correction of the varietal name usually cited as M.
anisopliae var. major.
89
TABLE 1. Conidial measurements of Metarhizium flavoviride. Isolates
(listed by ARSEF numbers) were grown on Sabouraud dextrose agar + 1% yeast extract
at room temperature. Sizes expressed as mean + standard deviation for 75 conidia
measured in lactophenol/aniline blue.
ARSEF HOST/SOURCE COUNTRY : SIZE (um)
M. flavoviride var. flavoviride
21334 — Ceutorrhynchus macula-alba Czechoslovakia 7-9(-11) x 4.5-5.5>
1184 Otiorrhynchus sulcatus France Seats i leks 3.080.)
2025 agricultural soil Fed. Rep. Germany 8.7+2.9 x 4.5+ 0.6
2026 agricultural soil Netherlands S20.9 x7 5.4 2 0.6
M. flavoviride var. minus b
2037" Nilaparvata lugens Philippines 4.6+0.7 x 2.7+0.3
1099 Nilaparvata lugens Philippines 46+0.6 x 2.3+0.4
1547 Recilia dorsalis Philippines 5.2+0.8 x 2.6+0.4
1764 Nilaparvata lugens Solomon Islands 5.4+0.7 x 2.6+0.4
1768 Nilaparvata lugens Solomon Islands 5.9+0.8 x 2.5+0.4
2023 Acridid grasshopper Galapagos Islands 4.9+0.7 x 2.6+0.4
4 Type strain of variety.
From Gams & Rozsypal (1973); the type strain subculture examined here, as noted by
Gams and Rozsypal, was unpigmented and produced few typical phialides but many
extremely variable clavate to cylindrical conidia, 8.6 + 1.7 x3.0+ 0.5 um.
variety of this species; these isolates consistently produce smaller conidia
and sometimes form fasciculate masses of hyphae (synnemata) in artificial
culture (Fig. 2b). Varieties of this species are characterized primarily by
conidial dimensions.
Metarhizium flavoviride Gams & Rozsypal var. flavoviride, Acta
Bot. Neerl. 22, 518-521 (1973). FIG. 1A
Cultures Examined: ARSEF 2133 (ATCC 32969 <CBS 218.56), ex type
culture, from larvae/pupae of Ceutorrhynchus macula-alba, Brno, Czechoslovakia,
1956. ARSEF 1184 (INRA MF-88 = CBS 700.74), from Otiorrhynchus sulcatus,
Brittany (France), January 1974. ARSEF 2025 (CBS 125.65), from soil under
Brassica oleracea, Kiel-Kitzeberg, Fed. Rep. Germany, 1963 or 1964. CBS 380.73,
from soil under Brassica oleracea, Kiel-Kitzeberg, Fed. Rep. Germany, 1963 or
1964. ARSEF 2026 (CBS 473.73), from agricultural soil, coll. J. W. Veenbaas-
Rijks, Wageningen, Netherlands, 1973.
Metarhizium flavoviride Gams & Rozsypal var. minus Rombach,
Humber, & Roberts, var. nov. FIGS. 1B, 2
Morphologiae conidiophororum, phialidium clavatarum, conidiorum ellipsoideorum
ovoideorum, et habitu in cultura Metarhizium flavoviride simulant, autem conidia
viridi-grisea parviora, 4.5-7 x 2-3 tm, et (in cultura) synnemata albida laxe fasciculata
sparsim fecunda formans.
90
Holotype: CUP 61454, on Nilaparvata lugens Stal, experimental rice
field, International Rice Research Institute, Los Banos, Philippines,
leg. M.C. Rombach, 5 November 1985.
Isotypes: CBS, IMI, and at the USDA-ARS Plant Protection Research
Unit (Ithaca, New York).
Cultures Examined: ARSEF 2037, type culture isol. from holotype, leg. M.C.
Rombach, November 1985. ARSEF 543, from greenhouse culture of Recilia
dorsalis (Motschulsky), International Rice Research Institute, Low Bafios, Laguna,
Philippines, D. W. Roberts, August 1980. ARSEF 1547, from Recilia dorsalis
(Motschulsky), Pangasinan, Luzon, Philippines, leg. R. M. Aguda, September
1982. ARSEF 1763 through 1773 (inclusive), from Nilaparvata lugen s Stal on
rice, Guadalcanal Plains, Guadalcanal, Rep. Solomon Islands, coll. by D. T. Ho,
isol. by M. C. Rombach, February 1985. ARSEF 2023 (CBS 544.81, as M.
flavoviride) from Acrididae (Orthoptera), Santa Fé Island, Galapagos Islands, coll.
H. C. Evans, 1981. Numerous cultures from Nilaparvata lugens on rice from the
International Rice Research Institute, Los Bafios, Philippines.
The branched conidiophores, clavate conidiogenous cells, and long
chains of ellipsoidal conidia of M. flavoviride var. minus generally
resemble those of M. flavoviride Gams & Rozsypal (1973) but differ in
their dimensions. The conidial chains of both varieties aggregate into
columnar masses similar to those of M. anisopliae. The conidiogenous
cells of var. minus are 8.4+ 1.2 x 2.8+0.3 um (on Sabouraud dextrose
5 um
Cy Oats).
0000. 7) COG
Fig. 1. Conidiophore and conidia of (a) Metarhizium flavoviride var. flavoviride
(isolate CBS 380.73a), (b) M. flavoviride var. minus on the brown planthopper, and (c)
in culture, ARSEF 1547.
of!
agar + 1% yeast extract) whereas those of var. flavoviride are 9-14 x
3—4.5 um (from malt extract and oatmeal agars). The conidia of var.
minus are 4.5-7 x 2-3 um and dull grey-green in mass (with no significant
hint of yellow pigments); the larger (6.5-11 x 3.5-5 um) conidia of var.
flavoviride are usually yellow-green. Conidial sizes in several strains of
these varieties are listed in Table 1. Because conidia of both barieties tend
to become slightly narrower after repeated transfers on artificial media;
isolates should be frozen in liquid nitrogen or lyophilized at the earliest
chance. In culture, M. flavoviride var. minus may produce synnemata up
to 10 mm high (Fig. 2b) rather than the more usual flat, stroma-like
sporodochia. These synnemata often remain white and sterile or may
show fertile patches.
Metarhizium flavoviride var. minus resembles M. album Petch in the
morphologies of the conidiophores, clavate phialides, and conidia. These
species differ in their conidial colors since M. album is whitish-brown
(with no hint of green pigmentation). Also, the flat, smooth hymenial
surface with conidial chains adhering in large columns produced by
M flavoviride var. minus differs markedly from the raised sporodochial
masses of hyphal bodies produced by M. album. The recognized species
of Metarhizium are also distinguished unambiguously by comparisons of
the electrophoretic mobility of their various isoenzyme systems.
fp
Fig. 2. M. flavoviride var. minus. (a) Infecting brown planthopper; insect on right
is an early stage of postmortem development with protruding white mycelium;
specimen on the left exhibits a darker fertile patch. (b) Synnemata of ARSEF 1547 in
culture; note apical fertile patches (fp). Both micrographs, x 7.
92
To date, the small-spored variety of M. flavoviride has been collected
from plant- and leafhoppers on rice in the Philippines and Solomon
Islands; it was absent from extensive collections of fungi from auchenor -
rhynchid hosts from rice in South Korea, India, and Sri Lanka. A
collection of Metarhizium flavoviride on a grasshopper from the
Galapagos Islands (Evans & Samson 1982) also belongs to var. minus
(see Table 1). Clearly, this variety may prove to be a much more widely
distributed and important pathogen of insects than 1s now apparent.
Acknowledgments
We thank Dr. B.M. Shepard for his continuous encouragement; Maureen
Rombach for preparing the drawings; the Curators at the CBS and ATCC
for providing cultures; Dr. D.T. Ho for the Solomon Islands collections;
Drs. W. Gams, W.B. Kendrick, R.P. Korf, and K.A. Seifert for their
constructive reviews: and the Office of the Science Advisor of the US
Agency for International Development for partly supporting this work.
References
EVANS, H.C., & R. A. SAMSON. 1982. Entomogenous fungi from the Galapagos
Islands. Can. J. Bot. 60: 2325-2333.
GAMS, W., & J. ROZSYPAL. 1973. Metarhizium flavoviride n.sp. isolated from
insects and from soil. Acta Bot. Neerl. 22: 518-521.
MARCHAL, M. 1977. Fungi Imperfecti isolés d'une population naturelle
d'Otiorrhynchus sulcatus Fabr. (Col.; Curculionidae). Rev. Zool. Agric. Pathol.
Végét. 76: 101-108.
POPRAWSKI, T. J..M. MARCHAL, & P. H. ROBERT. 1985. Comparative
susceptibility of Otiorrhynchus sulcatus and Sitona lineatus (Coleoptera: Curculi -
onidae) early stages to five entomopathogenic Hyphomycetes. Environ. Entomol.
14: 247-252.
POPRAWSKI, T. J., P. H. ROBERT, I. MAJCHROWICZ, & B. BOIVIN. 1985.
Susceptibility of Delia antiqua (Diptera: Anthomyiidae) to eleven isolates of
entomopathogenic Hyphomycetes. Environ. Entomol. 14: 557-561.
ROMBACH, M. C., AGUDA, R. M., SHEPARD, B. M., & ROBERTS, D. W. 1986.
Infection of the rice brown planthopper, Nilaparvata lugens (Homoptera;
Delphacidae) by field applications of entomopathogenic Hyphomycetes (Deutero -
mycotina. Environ. Entomol.: in press.
ROMBACH, M. C., HUMBER, R. A. & EVANS, H. C. 1986. Metarhizium album
Petch, a pathogen of plant- and leafhoppers of rice. [In prep. for Trans. Brit. Mycol.
Soc.]
SOARES, G. G., M. MARCHAL, & P. FERRON. 1983. Susceptibility of Otior -
rhynchus sulcatus (Coleoptera: Curculionidae) larvae to Metarhizium anisopliae
(Deuteromycotina: Hyphomycetes) at two different temperatures. Environ.
Entomol. 12: 1886-1890.
TULLOCH, M. 1976. The genus Metarhizium. Trans. Brit. Mycol. Soc. 66: 407-
411.
MYCOTAXON
VOlw ANVIL Y pp. 95-98 October-December 1986
STUDIES ON CHINESE ASCOMYCETES.
3. ASTROSPHAERIELLA LAGENIFORMIS.
% %H
JING-ZHU YUE & OVE ERIKSSON
ABSTRACT
Rhynchostoma Karsten is discussed. The cor-
rect name for the single species recorded
from China, R. lageniforme Teng, is Astro-
sphaeriella lageniformis (Teng) J.-z. Yue &
O. Eriksson.
INTRODUCTION
About 20 species have been described in Rhynchostoma
Karsten, and at least five other species have been trans-
ferred to the genus from other genera. Some species have
been excluded, but the genus is still in need of revision.
It is currently classified in the Trichosphaeriaceae, but
the type species, R. minutum Karsten, is certainly not
closely related to Trichosphaeria Fuckel. The only species
of Rhynchostoma that has been described from China is R.
lageniforme Teng. We have examined original material of
this species and compared it with original material of the
generic type and some other species in the genus. We found
that, in fact, the Chinese fungus has bitunicate asci and
belongs in Astrosphaeriella H. Sydow & Sydow.
MATERIALS AND METHODS
For details about methods used, see Eriksson & Yue (1985)
and Yue & Eriksson (1985).
* Institute of Microbiology, Academia Sinica, Beijing,
China. ** Institute of Ecological Botany, University of
Umea, S-90187 Umea, Sweden.
94
RESULTS
Astrosphaeriella lageniformis (Teng) J.-z. Yue & O. Eriks-
son comb. nov.
Bas.: Rhynchostoma lageniforme Teng, Sinensia 7: 508 (1936)
Ascomata scattered - densely aggregated, erumpent and
finally almost completely free and with only the flat base
lower than the surface of the host, hemispherical or sub-
conical, with uneven surface, ca. 300-500 pm across, with
a long, cylindrical or conical beak; ascomal wall ca. 100
pm thick, carbonized, opaque.
Centrum hollow in dry ascomatas; hamathecium of anasto-
mosing paraphysoids, ca. 1 um wide.
Asci numerous, ca. 90 x 14 pm, cylindric clavate,
rather short-stalked, bitunicate, with thin ectotunica and
rather thin endotunica, without ring structures, 8-spored.
Ascospores (Fig. 1B) ca. 35-40 x 6-7 pm, narrowly fusi-
form - clavate, somewhat inequilateral, medially 1-septate,
upper hemispore slightly broader, hyaline at first, then
pale brown, smooth, without distinct perispore.
Coll.: China, Hainan Prov., Yen-sheng, on Cocos nucifera,
15.11.1934,” leg. 'S.Q. Deng ‘ns “921, det.. S.C... Tene i(CuF=
Fig. 1. Astrosphaeriella lageniformis.
(A). Ascomata. (B). Ascospores; hyaline,
immature spore (left) and pale _ brown, B
mature spore (right). - Coll.: CUP-CH177.
Magn.: (A). x22. (B). scale = 10 pm. Lasensaenal
a5
DISCUSSION
Karsten (1870) distributed Rhynchostoma minutum n. gen. et
sp. in his exsiccatum Fungi Fenn. Exs., n. 950. The short
generic diagnosis in the lable reads "Perithecia rostrata;
sporae fuscae, uniseptate". We have examined the exsiccatum
preserved in Helsinki (H). It consists of two small pieces
of old wood of Juniperus communis with numerous, scattered
ascomata of R. minutum. They are small, globose, immersed -
almost free. The beak is broken at the base in almost all
ascomata in this collection, only in a few it is broken at
the middle. We saw only one ascoma with an unbroken beak.
The ascomata (incl. the beak) appear black under the dis-
secting microscope.
Karsten (1890: 178) described another species with
larger ascomata, R. rubrocinctum. Petrak and H. Sydow
(1923: 375) have given a detailed description of this
species. We have studied semithin sections of an ascoma
from a Swedish collection (Torne Lappmark, Abisko fpar.,
Palnoviken, on old wood, 15.VI.1984, n. 84940, leg. V.
Alstrup, det. 0O.E., Herb. C), closely agreeing with Petrak
& H. Sydow’s description of the species, except that they
describe the colour of the beak as "aussen dunkel rotbraun,
innen, besonders oben, intensiv rot gefarbt". The ascomata
which we have seen (several collections) have apically
Fig. 2. Rhynchostoma minutum. Ascomata on old, naked wood.
Coll.: Alstrup n. 84940 (Herb. C). Magn.: x22.
96
swollen beaks which are usually bright red just below the
tip (Fig. 2). The colour is intensive, almost resembling a
cigarette glow.
Miller & von Arx (1962: 608) examined original material
of R. minutum as well as R. rubrocinctum and came to the
conelusion that the species are identical. They did not
mention either the strong red colour and or the surface
structure of the ascospores. Several investigators have
independently discovered that the spores are provided with
spirally arranged ridges (Jing-zhu Yue, Leif Tibell, Geir
Mathiassen). Mathiassen (1985: Pl. VId) published a SEM
graph of spores from a specimen with beak coloured red in
its apical part. We have SEM-graphs of spores from the
above mentioned collection (Alstrup, n. 84940) and from the
original material of R. minutum. The ascospores are very
similar (Fig. 3), and Mliller and von Arx are most probably
right that the two species are identical, and that the
correct name for the species is R.- minutum Karsten. There
seems to be a seasonal variation, grading from "minutum
type" to "rubrocinctum type" of fruit bodies (L.E. Muhr, in
Tate)
The type species of Rhynchostoma, R. minutum, thus can
be characterized as a lignicolous pyrenomycete with globose
Fig. 3. Rhynchostoma minutum. (A). Ascospores and empty
asci on the surface of the swollen part of the beak. From a
specimen with large ascomata and red-coloured top of the
beak. (B). Ascospore from a specimen with small ascomata
and black beak. - Coll.: (A). Alstrup n. 84940 (Herb. C).
(B). Fungi Fenn. Exs., n. 950 (Herb. H). Magn.: (A). line =
10 pm. (B). line => 1 pm.
oy
perithecia provided with long, apically usually bright red
beaks, unitunicate asci with 8, 1-septate spores with
spirally arranged ridges. Also worth mentioning is the
presence of a hyaline amorphous crust around the beak
(thickest at the base), which has previously been observed
by Petrak & H. Sydow (1l.c.). The original description of
Rhynchostoma was too vague and most of the species that
have been referred to the genus are alien elements. Rhyn-
chostoma lageniforme Teng has hemispherical - subconical
ascomata with a beak and 1-septate ascospores, but there
are numerous important differences, for instance, the
ascomal wall is very dark-coloured. The centrum consists of
paraphysoids and bitunicate asci with elongate-fusiforn,
smooth ascospores, and it has to be transferred to the
genus Astrosphaeriella H. Sydow & Sydow in Melanommatales.
The genus Astrosphaeriella was monographed by Hawks-
worth (1981). He accepted 4 species, one of them, A.
aosimensis Hino & Katumoto, on leaf stalks of Livistona
(Arecaceae). Rhynchostoma lageniforme was described from
leaf stalks of Cocos nucifera (Arecaceae). The ascospores
are quite similar to those of A. aosimensis, but the
ascomata are covered by host epidermis in that species,
while erumpent in R. lageniforme. It is possible that the
two taxa are substrate forms of one and the same species,
the correct name of which would be A. lageniformis (Teng)
J.-Z. Yue & O. Eriksson. However, for the time being, we
suggest that they are kept separate.
*TIt is very remarkable that ascospores with almost exactly
the same surface structure and ontogeny are known in
Caliciales (Tibell, pers. comm.; see Tibell 1975: 76 ). It
is difficult to find a better example of how dangerous it
is to base a classification on one criterion only. In this
very case we know from other characters that Rhynchostoma
and the Caliciaceae are taxonomically far from each other.
ACKNOWLEDGEMENTS
The authors are grateful to Academia Sinica (Beijing), the
Swedish Institute (Stockholm), and the Natural Science
Research Council (Stockholm) for financial support of our
work. We thank Prof. Dr. D.L. Hawksworth, Mr. L.E. Muhr,
and Dr. L. Tibell for valuable information, Dr. L. Holm and
Dr. V. Shindler for reading the manuscript, and Dr. V.
Alstrup and the curators of CUP, H and UPS for the loan of
material.
98
LITERATURE CITED
ERIKSSON, 0. & YUE, J.-Z. 1985. Studies on Chinese asco-
mycetes. 1. Phaeosaccardinula dictyospora. - Mycotaxon
22: 269-280.
HAWKSWORTH, D.L. 1981. Astrosphaeriella Sydow, a misunder-
stood genus of melanommataceous pyrenomycetes. - Bot.
Js LINN SOG O22 50-59
HINO, I. & KATUMOTO, K. 1956. Notes on fungi from western
Japan. Cl) a= Bull. Prac. (Agric...) famaciicg 1 Uni ere
257-274.
KARSTEN, P.A. 1890. Fragmenta mycologica. XXX. - Hedwigia
29: 176-179.
MATHIASSEN, G. 1985. Pyrenomyceter (Ascomyceter) pa Salix i
Troms. - Inst. biol. geol., Univ. Troms¢.
MULLER, E. & ARX, J.A. VON 1962. Die Gattungen der didymo-
sporen Pyrenomyceten. - Beitr. Krypt.-Fl. Schweiz
VIit2) si 1=922.
PETRAK, F. & SYDOW, H. 1923. Kritisch-systematische Origi-
naluntersuchungen tiber Pyrenomyceten, Sphaeropsideen
und Melanconieen. I. - Ann. Mycol. 21: 349-384.
TENG, S.-Q., 1936. The mycoflora of China (2). - Sinensia
7: 490-527.
TIBELL, L. 1975. The Caliciales of boreal .North America.
- Acta Univ. Upsal. 21: 1-136.
YUE, J.-Z. & ERIKSSON, 0. 1985. Studies on Chinese asco-
mycetes. 2. Sinodidymella verrucosa. - Mycotaxon 24:
293-300.
TORE,
\
ne
AX TH al 5 te ke E (Rhynchostoma Karsten).
48 BH gl BY WE — BY Ay! — Bk We KA, (R. lageniforme
Teng) BY LAG ARA BA #048 (Astrosphaeriella la-
geniformis (Teng) J.-z. Yue & 0. Eriksson.
MYCOTAXON
VOLO, XXVELS pp. 99-106 October-December 1986
THERMOPHYMATOSPORA, A NEW THERMOTOLERANT
GENUS OF BASIDIOMYCETOUS HYPHOMYCETES
Shun-ichi UDAGAWA!, Takeyoshi AWAO?
and
Samir K. ABDULLAH?
1 National Institute of Hygienic Sciences, Kamiyoga
l-chome, Setagaya-ku, Tokyo 158, Japan
2 Central Research Laboratories, Ajinomoto Co., Inc.,
Suzuki-cho, Kawasaki-ku, Kawasaki-shi 210, Japan
3 Department of Biology, College of Science,
University of Basrah, Basrah, Iraq
SUMMARY
Thermophymatospora fibuligera anamorph gen. et
anamorph sp.nov., a basidiomycetous hyphomycete,
isolated from cultivated soil at date palm plan-
tation in Iraq is described and illustrated.
The fungus is characterized by large, brown,
globose, tuberculate conidia and the formation
of hyphae with clamp connections. The tempera-
ture range for its growth is between 17-48°C,
the optimum 35-40°C, no growth occurring at 50°C.
During investigations on thermophilic fungi isolated
from a variety of substrates in Iraq by the third author,
an unusual soil-borne hyphomycete assignable to a basidio-
mycete anamorph was encountered. The fungus produces
large, brown, thick-walled, tuberculate conidia, suggestive
superficially of Myceliophthora, but from which it is dis-
tinct by the formation of basidiomycete-like hyphae with
clamp connections at the septa. As it cannot be accommo-
dated in any basidiomycetous anamorph currently known, the
new genus Thermophymatospora is here proposed for it. The
Specimen studied is preserved at the Mycological Herbarium,
100
National Institute of Hygienic Sciences, Tokyo (NHL). The
living culture has been placed in the American Type Culture
Collection.
Thermophymatospora Udagawa, Awao et Abdullah, anamorph
gen. nov.
Fungus thermophilus. Coloniae effusae, brunneae,
floccosae vel pulveraceae, plus minusve coactae. Hyphae
fertiles hyalinae vel subhyalinae, ramosae, laeves, sep-
tatae, fibulis amplis praeditae. Conidiophora micronemata.
Conidia holoblastica, acropleurogena super hyphis conidio-
genis oriunda, raro intercalaria, plerumque solitaria,
continua, flavo-brunnea vel brunnea, globosa vel subglo-
bosa, incrassata, tubercularia cum projecturis prominenti-
bus et costis infernis reticulatis ornata, a cellula coni-
diogena septo separata. Laccasis praesentes. Teleomor-
phosis versimile ad Basidiomycetem pertinens.
Species typica: Thermophymatospora fibuligera Udagawa,
Awao et Abdullah.
Etymology: thermo-, from the Greek for warm; phymato-
spora, from the Greek for warty spores.
Colonies spreading, brown, floccose or powdery, more
or less felty. Fertile hyphae hyaline to subhyaline,
branched, smooth-walled, septate, provided with clamp con-
nections. Conidiophores micronematous. Conidia holo-
blastic, borne terminally or laterally on conidiogenous
hyphae, rarely intercalary, mostly solitary, one-celled,
yellowish brown to brown, globose to subglobose, thick-
walled, with surface tuberculate with prominent projec-
tions interconnected by low straight ridges, separated
from the conidiogenous cell by formation of a crosswall.
Laccase present. Basidiomycete with unknown teleomorph.
Type species: Thermophymatospora fibuligera Udagawa,
Awao et Abdullah.
Thermophymatospora fibuligera Udagawa, Awao et Abdullah,
anamorph sp.nov. (Figs. 1-6)
Coloniae in agaro 'potato-dextrose' ad 37°C late
effusae, constantes ex mycelio basali coacto tenui et
conidio abundanti, superne floccosae vel subtomentosae,
primo albae, deinde brunneae vel avellaneae vel latericiae;
reversum brunneum vel valde callainum.
Hyphae fertiles hyalinae vel subhyalinae, ramosae,
tenues vel ex parte incrassae, laeves, 2-6 um crassae,
septatae, ad septum 1-2 fibulis amplis praeditae. Conidio-
phora micronemata. Hyphae conidiogenae terminales vel
laterales, hyalinae, rectae vel flexuosae, simplices vel
101
irregulariter ramosae, plerumque 10-35 x 2-4 um, tenues,
glabrae. Conidia holoblastica, acropleurogena super hyphis
conidiogenis oriunda, raro intercalaria, plerumque solita-
ria, globosa vel subglobosa, (18-)20-24(-25) um diam (sine
tuberculo), primo hyalina, deinde flavo-brunnea vel medi-
brunnea, incrassata, saepe valde tubercularia cum projec-
turis usque 1.5-3 um longis et costis infernis reticulatis
ornata, interdum cum vestigio basali et conspicue promi-
nentia apicali, a cellula conidiogena septo 3-4(-5) um lato
separata, facile germinantibus.
Holotypus: cultura NHL 2959, ex solo sativo isolatus
eSic,, sdragiahark, Basrah. Iraq,:in. 25.x1.1984,. legit.
S.K. Abdullah. In collectione fungorum ‘National Institute
of Hygienic Sciences (NHL), Tokyo, Japan'.
Etymology:fibuliger, from the Latin for clamp-bearing,
referring to the formation of clamps.
Colonies on potato-dextrose agar at 37°C spreading
broadly, attaining a diameter of 8.5 cm within 7 days,
consisting of a very thin basal felt with floccose to sub-
tomentose overgrowth, at first white, becoming brown
(Methuen 7E4-5, Kornerup and Wanscher, 1978) or Hazel to
Dark Brick (Rayner, 1970) from massed conidia; reverse
brown to dark turquoise (Methuen 7£5-24F4-5) .Discoloration
with 4% or 10% KOH negative. Reactions with a-naphthol and
guaiacol positive. Reaction with p-cresol negative. Reac-
tion with hydrogen peroxide + pyrogallol negative.
Colonies on oatmeal agar growing as on potato-dextrose
agar but less floccose, rather powdery, bearing abundant
conidia at center, brown (Methuen 6E6) or Cinnamon to Dark
Brick (Rayner, 1970); exudate Fulvous (Rayner, 1970);
reverse grayish orange to grayish brown (Methuen 6B3-8F3)
or Fuscous Black (Rayner, 1970).
Fertile hyphae hyaline to subhyaline, branched, thin
or partially thick, smooth-walled, 2-6 um wide, regularly
provided with 1-2 round clamp connections at each septum.
Conidiophores undistinguished from conidiogenous hyphae.
Conidiogenous hyphae borne terminally or laterally as side
branches on the fertile hyphae, hyaline, straight or flex-
uous, Simple or irregularly branched, usually 10-35 x 2-4
um, thin, smooth-walled. Conidia holoblastic, borne ter-
minally or laterally, rarely intercalarily on the conidio-
genous hyphae, mostly solitary, globose to subglobose,
(18-)20-24(-25) um in diam (without tuberculations), ini-
tially hyaline, becoming yellowish brown to mid-brown in
age, thick-walled, with surface often markedly tuberculate
with projections reaching a maximum length of 1.5-3 ym and
interconnected by low straight ridges in reticulate appear-
102
Fig. 1. Thermophymatospora fibuligera.
A. Fertile hyphae with clamp connections. B. Conidio-
genous hyphae and young conidia. C. Matured conidia.
103
ance, sometimes with a basal remnant and a prominent apical
elongation, separated from the conidiogenous cell by forma-
tion of a crosswall, 3-4(-5) um wide at the base, germi-
nating easily by mean of a single germ tube.
Temperature relations: thermotolerant with optimal
growth and conidial formation at 35-40°C (Table 1).
Specimen examined: ex soil at date palm plantation,
Saraji Park, Basrah City, Iraq, November 25, 1984,
isolated by S. K. Abdullah, NHL 2959 (Holotype).
Table 1. Radial growth and conidia formation of
Thermophymatospora fibuligera strain
NHL 2959 at various temperatures
Temperature Growth, PDA Conidia, PDA
(20) Ad 10d Ad 10d
17 0 1-2 -
20 2-3 15-18 - -
25 16-17 70-75 ~ at
30 46-50 >85 + +++
35 69-72 >85 ate Lay
40 78-80 >85 Las a+
45 13-15 70-73 - +
50 0 0
call 250 ba ORE Bile RAN DDE SNE Sida ear a SARS tna en Ya
Growth: colony diameter (mm) after 4 and 10 days.
Conidia: ++++, very abundantly produced; +++, abundant;
++, moderate; +, slight; and -, not produced.
Some members of Myceliophthora OY Chrysosporium are
macroscopically very similar to this new fungus, differing
clearly by the absence of clamp connections at the septa
of hyphae and by their association with ascomycetous teleo-
morphs (van Oorschot, 1980).
The taxonomic position of most thermophilic fungi has
been reviewed comprehensively (Cooney and Emerson, 1964;
Crisan, 1973; Samson and Tansey, 1977), and reveals that
very few basidiomycetous fungi are thermophilic or even
thermotolerant. Coprinus delicatulus Apinis (considered
to be the same as C. cinereus (Schaeff. ex Fr.) S. F. Gray)
and Phanerochaete chrysosporium Burdsall (anam.sporotrichum
pruinosum Gilman et Abbott) grow at temperatures over 40°C.
From the new fungus the thermotolerant hyphomycete s. prui-
nosum (often noted as S. pulverulentum Novobranova or ana-
morph of P. chrysosporium) can be easily distinguished by
the white to cream-colored colony, distinctly branched
conidiophores, smooth-walled conidia and the absence of
and
Tons
x400.
fibuligera.
th clamp connect
ae wi
ja.
d
Matured coni
A
1100.
x
dia.
i
aN
aN ‘ : \ 7 )
_ " 2 : S Z ANE \ ay : ae i
Thermophymatospora
2-6.
iogenous
d
igs.
idi
F
2-4
young coni
6
hyph
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Con
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of matured con
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E
S
105
clamps in its hyphae. Cardinal temperatures for growth of
S. pruinosum are as follows: minimum 7°C, optimum 36-40°C,
and maximum 46-49°C (Stalpers, 1984). Therefore, both
fungi exhibit a marked difference in the tolerable limit
at the low temperature. Disporotrichum dimorphosporum
(von Arx) Stalpers (1984), another sporotrichum-1like
fungus, differs from the new fungus in having dimorphic,
smooth-walled, pinkish or hyaline conidia and the occa-
sional presence of multiple clamps in its hyphae. The
temperature optimum for growth of D. dimorphosporum has
been given as 30-32°C and the maximum as 40°C.
Sporotrichum azureum Wright et von Arx (non s.azureum
Link) is characterized by the formation of hyphae with
rounded clamp connections and obovoid to pyriform, smooth
or irregularly verrucose conidia (von Arx, 1973). Accord-
ing to Stalpers (1984), s. azureum Wright et von Arx is
closer to Ptychogaster than to the three re-defined mem-
bers Of Sporotrichum. Aside from its thermophilism, the
unique ornamentation of conidia, which somewhat resembles
the wall structure of Ravenelia spegazziniana uredinio-
spores (Gardner and Hodges, 1985), serves to distinguish
the new fungus from Ptychogaster and related genera.
ACKNOWLEDGMENTS
We wish to thank Prof. R.T. Hanlin of the University
of Georgia for reviewing the manuscript; and Dr. Y. Otani
for correcting the Latin diagnosis.
LITERATURE CITED
Arx, J.A. von. 1973. Further observations on Sporotrichum
and some similar fungi. Persoonia 7: 127-130.
Cooney, D.G., and R. Emerson. 1964. Thermophilic Fungi,
an Account of Their Biology, Activities and Classi-
fication. Freeman, San Francisco. 188p.
Crisan, E.V. 1973. Current concepts of thermophilism and
the thermophilic fungi. Mycologia 65: 1171-1198.
Gardner, D.E., and C.S. Hodges, Jr. 1985. Spore surface
morphology of Hawaiian Acacia rust fungi. Mycologia
77: 575-586.
Kornerup, A., and J.H. Wanscher. 1978. Methuen Handbook
of Colour. 3rd ed. E. Methuen, London.
Oorschot, C.A.N. van. 1980. A revision of Chrysosporium
and allied genera. Stud. Mycol. 20: 1-89.
Rayner, R.W. 1970. A Mycological Colour Chart. Common-
wealth Mycological Institute, Kew, Surrey & British
Mycological Society.
106
Samson, R.A., and M.R. Tansey. 1977. Guide of thermo-
philic and thermotolerant fungi. Abstracts, Second
International Mycological Congress, Tampa, Florida.
5p.
Stalpers, J.A. 1984. A revision of the genus Sporotrichum.
Stud. Mycol. 24: 1-105.
MYCOTAXON
ese
VOLE AXVAd .upparlO7s111 October-December 1986
eee eee
GIBBAGO, A NEW PHAEODICTYOCONIDIAL GENUS OF HYPHOMYCETES
Emory G. Simmons
Department of Botany, University of Massachusetts
Amherst, Massachusetts 01003
Gibbago trianthemae Simmons, sp. nov., type of a new
genus, is described from parasitized leaves of Trianthema
portulacastrum L. (Aizoaceae) collected in Cuba, the USA
(Texas), and Venezuela. Genus characters are compared with
those of somewhat similar genera Alternaria, Embellisia,
Stemphylium, and Ulocladium.
GIBBAGO Simmons, gen. nov. Rigi.
Etym.: gibbus (hump, swelling) + -ago (indicating
presence of a property), with reference to swollen
conidial cells that function as conidiophores.
Conidiophora ex hyphis lateraliter oriunda, solitaria
vel 2-4-fasciculata, simplicia vel raro ramosa, recta vel
in locis sporiferis geniculata, septata, brunneola. Coni-
dia initio solitaria, ellipsoidea, erostrata, leviter
colorata, septis transversalibus longitudinalibusque,
cellulis apicalibus tumentibus et conidia secondaria effe-
rentibus. Typus:
Gibbago trianthemae Simmons, sp. nov. Conidiophora
simplicia vel 1-2 geniculate conidiogena, straminea, usque
ad 60-80 x 5-6ym. Conidia primigena solitaria, pellucide
straminea, laevia, ellipsoidea, demum late ellipsoidea vel
late subovate ellipsoidea, vulgo 35-45 x 15-22ym, plerumque
1-4 transverse et 2 longitudine septata, ad mediseptum
transversum leniter constricta, poro basali inconspicuo;
cellulis apicalis 1-4 tumentibus et conidia secondaria
isdem ac conidia primigena efferentibus. Habitatio typi in
laesionibus foliorum Trianthemum portulacastrum L. Typus:
EGS 21-051, E. G. Simmons, 27-VIII-1967, College Station,
TX, USA, lectus; partes in BPI (holotypus) et pro isotypis
saltem in IMI et NY conservandae.
108
GIBBAGO Simmons, gen. nov. Conidiophores solitary or
2-4 loosely fasciculate, erect, rarely distantly branched,
simple with a single apical conidiogenous locus, then often
proliferating by means of a secondary conidiophore that
arises immediately below the apical cell of the existing
conidiophore; septate, slightly pigmented. Conidia initi-
ally solitary, ellipsoid, beakless, pigmented, becoming
transversely and longitudinally septate; with apical cells
swelling slightly and producing secondary conidia similar
to initial ones. Type:
Gibbago trianthemae Simmons, sp. nov. Conidiophores
simple or 1-2 proliferated, 1-4-transeptate, pale straw-
colored, up to 60-80 x 5-6ym, very slightly swollen at
apex, producing a solitary conidium at the apex of each
proliferation, retaining a distinct umbilicate or crateri-
form depression at the conidiogenous locus after secession
of conidium. Conidia initially solitary, almost perfectly
ellipsoid; becoming broadly ellipsoid to broadly subovate-
ellipsoid, with 1-4 complete or partial transverse septa
(slightly constricted at initial median septum), 2 complete
longitudinal septa intersecting at right angles in each
conidium half, plus a few shorter ones in transverse sec-
tors of the conidium; clear pale yellow-brown, smooth; with
a minute basal pore-scar that is difficult to observe and
that lacks any sort of complex surrounding structure or
halo of pigmentation; commonly maturing at about 35-45 x
15-22,m; with 1-4 of the apical cells enlarging slightly
and each giving rise directly to a single secondary coni-
dium morphologically identical with primary conidia; indi-
vidual apical (sometimes basal) cells also sometimes giving
rise to conidiophores that have the distinctive apically
swollen and umbilicate appearance of hyphal conidiophores
and that are about 7 x 6um.
Type EGS 21-051, on well-defined, often extensive
lesions in leaves of Trianthema portulacastrum L., coll.
E. G. Simmons, Texas A. & M. University Experiment Farm,
College Station, TX, 27 Aug. 1967; dep. BPI (holotype) and
also at least in IMI and NY (isotypes).
All of the nine collections of this fungus known to me
are on horse-purslane, a common pantropical weed of culti-
vated fields. The earliest collection (BPI: Runyon 4480,
Sept. 1953) is from Texas, as are a recent isolate con-
tributed by J. K. Mitchell (EGS 38-128) and my own collec-
tions (EGS 21-051, 21-052); collections in IMI (coll. R.
Urtiaga, 1966-1969) are from Cuba and Venezuela.
Fig. 1. Gibbago trianthemae, conidia and conidiophores;
conidium at upper left exhibits two apical cells that have
sporulated directly and two that have produced hyphal
conidiophores. Vertical bar = 50yn.
110
Gibbago trianthemae can be established readily in
culture by means of 1l-conidium isolates. It grows well and
sporulates abundantly in plates of PCA, Hay, and V-8 agars
held at moderate temperatures under cycled light (cool-
white fluorescent, light/dark, 8/16 hrs). Subsurface
mycelial growth is dense and dark on V-8, less so on PCA,
and inconspicuous on Hay. Sporulation is excellent at agar
surfaces and on the moderate amounts of medium gray, woolly
aerial myceliun. Conidia produced in culture are more
likely to produce secondary conidiophores and conidia than
they are to sporulate precociously directly from their own
apical cells. Primary conidiophores produced in culture
commonly are 200-300ym long, secondary ones 60-70vuym.
Representative 1l-conidium isolates ex Type include EGS
21-046 through 21-050; EGS 38-128, received in culture from
J. K. Mitchell, is a typical strain. Some of these repre-
sentatives are being deposited in international culture
collections.
Discussion
For many years I have juggled this intriguing fungus
among species of several established genera of phaeodictyo-
conidial Hyphomycetes (Alternaria, Embellisia, Stemphylium,
Ulocladium), but without enough conviction to publish it as
novel in any one of them.
Cfr. Stemphylium: Gibbago trianthemae conidia, before
they proliferate, are stemphylioid in general appearance
except that they lack the pigmented halo and the umbilicate
depression that usually are conspicuous around the basal
detachment scar of Stemphylium conidia; conidiophores of
the species and their manner of proliferation are not at
all those of Stemphylium, whose conidiogenous cell charac-
teristically has a subapical pigmented halo and which
proliferates by means of a secondary conidiophore that
originates in or near the basal septum of a preceding
conidiogenous cell and then emerges through the apex of
that cell.
Cfr. Alternaria: conidia of G. trianthemae might be
considered erostrate alternarioid, of the sort known for A.
helianthi and A. leucanthemi, but here again the lack of a
basal, pigmented halo and conspicuous detachment scar gives
pause; conidiophores have the kind of laterally geniculate
proliferation characteristic of Alternaria.
Cfr. Ulocladium: conidia of G. trianthemae have some
characteristics in common with those of Ulocladium, in that
the detachment scars of both are very inconspicuous, lack a
(rg
pigmented halo, and develop on laterally geniculate coni-
diophores (as in Alternaria). However, G. trianthemae
conidia are not initiated as distinctly obovate, basally
pointed juveniles, which are generically typical of Ulocla-
dium.
Cfr. Embellisia, which is distinguished to a major
extent by its species' rigid, wide-band, transverse coni-
dial septa and the semi-collapsing nature of the outer
conidial wall, both of which characters are foreign to
Gibbago as well as to the other genera discussed.
A genus would be established on shaky ground, of
course, if it were defined only by such an accumulation of
small negative observations. In characterizing Gibbago, a
unique pattern of secondary sporulation distinguishes the
genus from all other phaeodictyoconidial Hyphomycetes with
some otherwise similar characters.
Most notably, the rounded distal end of a mature G.
trianthemae conidium has four cells delimited in part by
the right-angle intersection of two longitudinal septa.
The apex of each of these cells may become slightly but
distinctly enlarged and then, without further change in
morphology or pigmentation, produce a secondary conidium;
only a minute pore remains at the locus of production after
a conidium secedes.
It must be emphasized, in view of the increasingly
common use of cultures in identification procedures, that
an individual cell in any part of a conidium (but usually
in the apex or the base) also may germinate and produce
simple conidiophores morphologically indistinguishable,
except for origin, from those formed on vegetative hyphae
(Fig. ob. upper deft)... These conidiophores may be short or
long (7-70ym), are somewhat swollen terminally, and exhibit
an umbilicate apex after release of a conidium. They are
germination hyphae converted to a special function; similar
ones are produced in most species of the several genera
discussed above. However, except for the act of conidium
production, they have no resemblance to the directly sporu-
lating cells of Gibbago trianthemae conidia, which are held
to be typical of the genus.
Ie Rel Ie
Margaret E. Barr Bigelow continues to be the subject
of my thanks for her prepublication editorial vetting of
this manuscript.
MYCOTAXON
VOL) SV EES opp ek Se) 7 October-December 1986
A NEW SPECIES OF THE LICHEN GENUS SULCARIA
(ASCOMYCOTINA, ALECTORIACEAE) FROM CALIFORNIA
IRWIN M. BRODO
National Museums of Canada, Museum of Natural Sciences,
Ottawa, Ontario K1lA OM8, Canada.
SUMMARY
A puzzling alectorioid lichen was
discovered in southern California that seems
best classified as a Sulcaria. It is caespitose,
abundantly isidiate, and contains protocetraric
acid. It is named here as a new species:
Sulcaria isidiifera Brodo.
Halfway between San Franscico and Los Angeles, in Los
Osos Oaks State Reserve, a very odd alectorioid lichen was
collected that is remarkable with respect to chemistry and
morphology. Although sought in numerous nearby areas in
California .by, the original collectors, Charis Bratt. and
Mason Hale, the lichen is so far known only from this one
forest. The locality is a very special one, with ancient
(600-800 year old) trunks of Quercus agrifolia forming a
climax community over stabilized sand dunes originally
covered by scrub and chaparral (Wieman 1980). It is also
the type locality of the recently described Hypogymnia
mollis Hale, a species possibly related to a Moroccan
taxon (Pike & Hale 1982).
This lichen proved extremely difficult to classify as
to genus. The genus Alectoria in the broad sense was
recognized to be assemblage of closely related genera by
Brodo & Hawksworth (1977), including Alectoria s. str.,
Bryoria and Sulcaria, distinguished by a combination of
chemical, anatomical, and especially spore characteris-
tics. This new species, immediately distinctive by virtue
of its longitudinally split, abundantly isidiate-sorediate
Dranches, and its"'“productiton "of> protocetraric acid, is
unfortunately without apothecia. Spore characters, which
would immediately serve to classify the species (2-4
brown, simple spores per ascus in Alectoria; 8 colorless,
Simple spores in Bryoria; (6-)8 colorless, 1(-3) septate
spores in Sulcaria), were therefore unavailable. Repeated
attempts by Bratt and Hale to find thalli with fruiting
bodies proved unsuccessful.
114
The branches of the new lichen are split longitudi-
nally as they are in species of,Sulcaria. The branches are
not! hollow! as | in Ss. sulcata', however, but are packed
with hyphae. In fact, the "sulci" are filled with soredia
from which clusters of short or long spinulose isidia
develop (Fig. 3). (The branches of S. virens and S.
badia, however, are not hollow either.) <
The production of protocetraric acid and (in one
specimen) a trace of what appears to be virensic acid,
links the species best with Sulcaria, where known species
produce psoromic or virensic acid, or atranorin. A
chemicai parallel can be found in the Lecanora
caesiorubella group, where these four compounds occur in
closely related species. Bryoria is also characterized
by the production of beta-orcinol depsidones, whereas no
species of Alectoria has these compounds. Although traces
of protocetraric acid occur ,in species ‘of! Bryorira
containing fumarprotocetraric acid as a principal
constituent, this substance has not been found as the main
product in any known Bryoria.
Anatomically, the new species most resembles’ an
Alectoria (with thick, knobby medullary hyphae and
decomposed cortical surface), or a Bryoria sect. Implexae
(its cortical hyphae having very little strengthening
matrix), but the cortical structure of Sulcaria badia and
S. virens is somewhat similar.
With respect to color, our material is closest to
Sulcaria anda few species of Bryoria sect. Implexae (such
as) By | Spirali tera, Or UB. kuemmerleana). No species of
Alectoria has the same badious tint.
The isidiate soralia seen in the new species are
rather unique in the Alectoriaceae. They develop from a
gradual disintegration of the cortex rather than froma
fissuring’ as in Bryoria furcellata (or 8B.) smithii,; orga
proliferation from a raised pseudocyphellum as in
Alectoria imshaugii. Viewing the various developmental
stages of sulca formation temporally, one can describe the
Situation as follows: At first, a few short isidia grow
superficially on a = branch. The cortex under and around
the isidia is rough and is obviously deteriorating. The
isidia develop along a "line" running down the branch. The
cortex finally disintegrates along this line, which
becomes transformed into a sorediate sulca filled with
spinulose isidia of various lengths. Sometimes the sulcae
are moderately deep, but most are simply wide and packed
with granulose soredia and isidia. The proposed name of
the new species is based on these unusual structures.
It would seem that the new species is closely allied
to either or both Sulcaria badia and Bryoria spiralifera.
eomencia ane follows Brodo & Hawksworth (1977).
115
Material of the latter species found in the Los Osos Oaks
State Reserve occasionally produces an incipient sulca
instead of a spiralled pseudocyphellum bringing it closer
to Sulcaria morphologically. In view of the fact that all
of these species are invariably sterile, I can see no
point in speculating on more appropriate generic designa-
tions at the moment. It does, however, make me more
inclined to classify the new species as a Sulcaria.
Sulcaria isidiifera Brodo, sp. nov. Figs. 1-3
Thallus caespitosus, cremens’ vel pallissime brunneus
vel badius, opacus; rami principales iso-dichotomiter vel
aniso-dichomiter fastigiati, ramis lateralibus spinulosis
plus minusve perpendicularibus, sulcati, sulcis isidio-
sorediatis spinuliferibusque. Apothecia et pycnidia non
vidi. Thallus acidum protocetraricum continens.
Typuee U.S.A.;, California, San Luis |Obispo ‘County: Los
Osos Oaks State Reserve, Los Osos Valley Road, alt. 100
ft., in Adenstoma fasciculatum. Bratt 3871, et J. Larson,
8 Jan. 1984 (holotypus: CANL; isotypi: US, hb. BRATT).
Thallus dull yellowish-white (927) grading into
light brown (57) and reddish brown (42) at the more
exposed tips; rarely shades of olive-grey (112) in places
(perhaps ecologically influenced); caespitose, 3-5 cm
long; main branching isotomic to anisotomic dichotomous,
with more or less perpendicular spinulose branches
developing from splits in the thallus; branches splitting
lengthwise and opening into rather wide, linear soralia
filled with spinulose isidia and spinules, often with
brown tips; main branches 0.3-0.5 mm wide; secondary
branches 0.15-0.3 mm wide; branches fairly even and smooth
except for the sulci and isidial development; branches
very brittle; pseudocyphellae apparently absent, or
developing immediately into isidiate soralia. Apothecia
and pycnidia not seen.
Copcexs) KOH=—9) C=) UoKC="" or) faint: pink, (PD+) “orange;
medulla KOH-, C-, KC-, PD-or faint orange. Contains
protocetraric acid, rarely with an unidentified compound
(R Classes 3, 5-6, 5: Culberson 1972) and a trace of
(2) virensic acid as accessory substances.
Habitat: On branches of Quercus agrifolia, Adenostoma
fasciculatum, and Coenothus ramulosus in sandy areas.
Distribution: Known only from the type locality.
Specimens seen: U.S.A. California. San Luis Obispo
County: Los Osos Oaks State Reserve, Bratt 3871 (type),
3851B, 4029, 3859 (hb. BRATT, CANL); Hale 57816 (CANL, US)
Numbers following colors refer to Kelly (1965).
a7,
ACKNOWLEDGEMENTS
I am indebted to Charis Bratt and Mason Hale in
bringing this interesting taxon to my attention, and for
trying so persistently to find fertile material. I also
thank Dr. Hale for revising the latin description, and
David Hawksworth for his views on the problem.
LITERATURE CITED
Brodo, I.M. & D.L.Hawksworth. 1977. Alectoria and allied
genera in North America. Opera Botanica 42: 1-164.
Culberson, C.F. 1972. Improved conditions and new data
for the identification of lichen products by a
standardized thin-layer chromatography method.
Journal of Chromatography 72: 113-125.
Kelly, K.L. 1965. ISCC-NBS color name charts illustrated
with centroid colors. -Supplement to national Bureau
Oprsorangards Circular s553.—"Washington, D.C.
Pike, L.H. & M.E. Hale, Jr. 1982. Three new species of
Hypogymnia from western North America (Lichenes:
Hypogymniaceae). Mycotaxon 16: 157-161.
Wieman, H. 1980. Nature walks on the San Luis coast.
Padre Publications, San Luis Obispo, California.
em ce es cs ee ee em es ee ee ee ee re ee ee ee ee ee ee ee ee ee ee
Figures 1-3. Sulcaria isidiifera, holotype (Bratt 3871).
-l. A single thallus. Scale = 5 mm. -2. Branches, showing
shallow sulci (arrow). Scale = 2 mm. -3. Close-up of
isidiate soralia and spinules. Scale = 1 mn.
MYCOTAXON
Moo assy pp. aL) 2 5 October-December 1986
BADARISAMA SOJAE, A NEW BULBIL-FORMING GENUS AND SPECIES
ny aye KUNWAR-, Va, Jae MANANDHAR*, and 308.05 CUNCLALR
Department of Plant Pathology, University of Illinois at
Urbana-Champaign, 1102 S. Goodwin Avenue, Urbana, IL 61801
Abstract
A hitherto undescribed bulbil-producing fungus, Badarisama
SOtdoucgen. peulsp.. NOV risydescnibed.. 7) tt has beenyobserved
frequently growing with Alternaria alternata from surface-
sterilized soybean) (Glycine) max); seeds on potato-dextrose
agar plates (25°C). Its colonies produce pseudoparenchyma-
tous, raspberry-like, black bulbils. No other method of
sporulation is known.
Introduction
A bulbil-producing fungus has been encountered growing in
association with Alternaria alternata (Fr.) Kiessler on soy-
Dean (Glycine max (l%).Méerr.) seeds (10). Weresub: & LeClair
(11) used the term ‘'bulbils' for papulospore-like structures
made up of pseudoparenchymatous tissue. Papulospores were
described by them as "thallodic propagules differentiated
almost from the inception into central and sheathing cells".
The term 'spore-knaueln' used by Eidam (2) for the papulo-
spores of Papulasporea parasiticum (Karsten) Harz was based
on a definition by Dodge (1). Fungi producing bulbils
rather than true papulospores, which had previously been in-
cluded in Papulaspora Pruess by Hotson (6,7), were removed
by Weresub & LeClair (11) to other genera. Bulbils have
been recognized only in the basidiomycetous genera Burgoa
Goid. and Minimedusa Weresub & LeClair (5). Our bulbil-
producing fungus differs from these genera.
Materials and methods
During 1984, symptomatic soybean seeds were selected at
random and surface sterilized for 5 min with 0.5% NaOCl (10%
Clorox) and then washed in two changes of sterilized, de-
ionized distilled water. The seeds were plated on 9-cm cul-
ture plates containing potato-dextrose agar (Difco) acidi-
fied (pH 4.5) with lactic acid (APDA) and incubated unde
12-hr photoperiods of cool fluorescent light (800 uEin/m’ /
sec) for 5 da at 25°C. We frequently observed and isolated
a bulbil-producing fungus consistently associated with
colonies of A. alternata.
Present. address, & An (¢. Kunwar, Raman Research Institute,
Bangalore 560080, India
Present address: Division of Plant Pathology, Department of
Agriculture, Lalitpur, Nepal.
120
To assist in the identification, in selecting a suitable
medium for growth, and to try to induce reproductive struc-
tures other than bulbils, the fungus was cultured on the
following media and incubated in the 12-hr photoperiod des-
cribed previously or in continuous dark or continuous light:
3.5% Czapek-dox agar, 2.3% lima bean agar, 3.5% malt-extract
agar, 7.3% oat-meal agar (OMA), nonacidified PDA, vegetable-
juice agar, and 2% water agar, each with or without two to
three autoclaved-sterile, 5-cm alfalfa (Medicago sativa L.)
or soybean stem pieces per plate, or halved soybean seeds,
or green pods (four to five pieces/plate) added aseptically.
To confirm that it was not a sclerotium-forming Alternaria
(3), individual bulbils and single Conidia of A. alteéeriara
were subcultured repeatedly on APDA and OMA and studied un-
der a binocular bright-field microscope. Living cultures
were compared with those of Burgoa verzuoliana Goid. (ATCC-
24040) and Papulospora pannosa Hotson (ATCC-24406).
A. ‘alternata, and, the bulbil-forming tungus were ystudiecaun
dual cultures. Five-mm discs were cut from the edge of pure
cultures of colonies of either fungus and both fungi placed
adjacent to one another at the center of each of four 9-cm
APDA culture plates and incubated under light as described
previously. Culture plates with either fungus alone served
as, controls.
Cultune filtraresiof As alternata; irom seultures “Growneps
2.4% potato-dextrose broth for 2 wk at 25°C, were filtered
through 022 um) pore Nalgene filters; and* mixed with) PDA:
20 v/v), and used to determine any effect on the growth of
the bulbil-forming fungus. Five-um discs were cut from the
edge of PDA colonies of the latter and placed in the center
of 9-cm culture plates amended with the culture filtrate of
A. alternata. Cultures grown on nonamended PDA served as
controls: *Growth) ratefwasorecordedi atten 7i daar. 207 Cc.
Taxonomic part
Our observations indicate that the bulbil-forming fungus
and A. alternata are discrete tandy unrelated taxa (Since mic
former appears to be unique in its morphology, it is herein
described as new.
Badarisama Kunwar, Manandhar et Sinclair. (Figs. 1-20)
(etym. Sanskrit - badari - berry, sama = like
Coloniae in agaro cum farina avenacea (OMA) vel cum dex-
troso et tuberibus Solani tuberosi (PDA) composito satae
immersae vel aliquid superficiales, effusae, fuscae; mycel-
ium ex hyphis ramosis, septatis, subhyalinis, zygodesmatibus
carentibus, maturis fusco-brunneis, compositum; bulbuli
blastici, e cellula vel hypha laterali ad angulum rectum ab
axe principali orti; bulbulorum cellulae crasse tunicatae,
pseudoparenchymaticae, homogeneae, brunneae, granulares, e
textura globosa in texturam angularem desinentes; bulbuli
instar fructuum Rubi, globosi vel subglobosi vel enormes,
Maturl fLuscw. ) Species typica: Be sojvae,
Colonies on OMA and PDA immersed to somewhat superficial,
effuse, dark; mycelium composed of branched, septate, sub-
Lat
hyaline hyphae lacking clamp connections, becoming dark
brown at maturity; bulbils blastic; arising laterally at
right angles to the main axis from a single lateral cell or
hyphae; cells of bulbils thick-walled, pseudoparenchymatous;
homogeneous, brown granular, composed of textura globulosa
to textura angularis; bulbils raspberry-like, globose to
subglobose or irregular, dark at maturity.
B. sojae Kunwar, Manandhar et Sinclair. CE VOSS ie it Os 20)
Coloniae in OMA satae 4.5-6 cm, in PDA usque ad 1.0 cm
diametro, in spatio septem dierum sub calore 25°C, atrae;
mycelium exiguum, procumbens, per vel super substratum
Figures 1-13. Badarisama sojae: Successive stages in the
development of a bulbil.
122
crescens, By En as UM cellulis multinucleatis, axe prinicip-
ar pulbilifero 4.2-7.2 um crasso; cellulae vel hyphae lat-
erales bulbiliferae 6.8-16.0 um crassae; bulbuli plerumque
in agaro immersi, fumosi, maturitate atri, forma magnitud-
ineque enormes, 91.0-218.4 x 136.0-236.6 um, medio 150.9 x
L71L.8 ‘um,, céllulis 97.0-18.5 x’ 16.4—-22.57 um) medio ls oe
Lo 6r Gms
Colony.diam 4.5 ‘to 6.0.icm on ‘OMA*and up to lcm. on (PDA
7 da at 25°C, black; mycelium scanty, procumbent, growing
on or in the substratum, hyphal cells multinucleate, bul-
biliferous main axis hyphae 4.2 to 7.2 um in width; bul-
biliferous lateral cells or hyphae 6.8 to 16 um in width;
bulbils mostly immersed in the agar, smoke-colored, becom-
ing bldck at maturity, variable in shape and outline, rang-
ing from,91.0' to 218.4 x 136.6) um; average Si1zeVio0 297s
171.8 cm, (cells an ‘the buibil ranging trom oie Vo. oe
16.4) to 22.35 um, average size Of vlo ao ox 7 oman
see
frie
ee
ey
Hee a
EE Nie
Figure 14. Badarisama sojae: A mature bulbil
Isolates examined: from soybean seed, Agronomy-Plant
Pathology South Farm, Univ. of Illinois at Urbana-Champaign
Urbana, IL,’ October) 1984, 1. K. Kunwar, ILLS-45312, holo=
type: and a living culture in the American Type Culture
Collection (ATCC-60345).
The colony diam of B. sojae was up to 1.5 cm in 7 da on
media other than OMA including APDA. When grown in dual
culture on APDA with A. alternata, colony diam was 6 to 7
cm, but 1 cm in the control. Mycoparasitism was not obser-
ved. When grown in PDA amended with a culture filtrate of
A. alternata, the rate of qrowthvor Bb. (so jae wasy2) tol je
cm.) (ite appears’ that A. alternate produces a factor or fac-
tors that stimulates faster and denser growth of B. sojae
InsecuLrires
Dao
Figures 15 to 20. Badarisama: 15. 7-da-old colony on
Oab-Meall agar wwscaltel=. li dvcm: tho to 100s iPhotomicrodqraphs
of stages in the development of a bulbil, scale = 15 um;
and 20. Photomicrograph of developing and a mature bulbil
on a hypha under bright-field microscope, scale = 62 um.
The mycelium was subhyaline, becoming thick-walled and
dark brownin3 to 4 da. Subhyaline hyphal cells contained
indefinite numbers of oil globules (Figs. 3-5, 16-18).
Septa were formed with a refractive thickened peripheral
ring. Anastomosis of hyphae was common. Clamp connections
124
were not observed.
Bulbils formed along the hyphae, blastic, developing lat-
erally ‘from ‘parts: of the. basal cell (Figs: 1-0, 16-18).
Cells within the bulbils develop successively in the same
manner from older mature cells. Ina few cases, bulbils
give rise to hyaline hyphae (Figs. 1-5, 7-11). Chains of
bulbils (are formed ‘in’ 2—- to. s-mo-old cultures (rigs... 4,
19% ZO
Discussion
Since 1B. ‘sojae’ reproduces ‘only by builbils;, we placed™t in
the Form-Order Agonomycetales (9). In Burgod,, cel tsaumsa
bulbil expand synchronously into individual bubbles. In
Minimedusa, bulbil primordia appear as a group of lateral
aerial hyphal branches converging to form a globose bulbil.
The dark bulbils of Badarisama can resemble either the dark
papulospores of Papulaspora atra Hotson and P. pannosa, or
dark bulbils of Burgoa nigra (Hotson) Goid. However,the
papulospore primordia of P. atra are formed from a group of
terminal cells on lateral branches (8), and mostly intercal-
ary an P. pannosa) (6,0)... ihe DuLbil is Obey. inigra arem uouned
by the synchronous expansion of bulbil cells (4). The bul-
bil/papulospore primordia and successive development of B.
versuoliana (ATCC-24040) and P. pannosa (ATCC-24406) were
different from Badarisama sojae. Therefore, Badarisama is
different from the bulbiliferous genera Burgoa and
Minimedusa by the presence of procumbent, immersed hyphae;
homogeneous, nonsynchronous and pseudoparenchymatous bulbils
formed from individual lateral cells or hyphae acting as
primordia; and the absence of clamp connections.
Acknowledgements
The authors wish*’to thank wJco eb Crane.) D..A Glawe aunt.
of Illinois at Urbana-Champaign: (UIUC), Urbana, LL) for
their valuable suggestions and advice, assisting in the
naming of the fungus, and for critical manuscript review;
to D. P. Rogers, Professor Emeritus, UIUC, for Latin diag-
nosis; and G. Morgan-Jones, Auburn Univ., Auburn, AL for
critical review of the manuscript.
Literature cited
1.) Dodge, Bs 0.) 19202. The vlifethtstory of Ascovolus
mangificus. Origin of the:.ascocarp £Lrom tEworstrains.
Mycologia, 1273115-134:
2. Eidam, E. 1883. Zur kenntniss der Entwicklung bei den
Ascomyceten. In Beitrage zur Biologie de Pflanzen. F.
Cohn. (ed 2 )mSi377—4 33.
3. Ellis, M. B. 1976. More Dematiaceous’ Hyphomycetes,.
Commonw. Mycol. Inst 750 /pp.
4. Goidanich, G. 1938. Studie sulla microflora fungina
cella pasta di legno destinata alla fabbricazione
cellarcanta, (Part d).)> Boll. (Stal Patol. Veo. f Roman...
17:305-399.
5... Hawksworth,/D. 1b.) 7SuttonsB: Cx, Ba Arnsworth? Greece
1983. Ainsworth and Bisby's Dictionary of the Fungi.
7th ed. Commonw. Mycol. Inst., Kew, Surrey, England.
like
125
Hotson, sH.,W: 1912. Culture studies ofl fungi producing
bulbils and similar propagative bodies. Proc. Amer.
Acad. Arts’ 6 SC. 48<277-306.
Hotson, H. W. 1917. Notes on bulbuliferous fungi with
a key to described species. Bot. Gaz. 64:265-284.
HOtsoOn, Hu. We 1929. PapulLospora attra nM... Sp. Amer. J’.
Bot.) 202219-220),
Kendrick, W. B. & Camichael, J. B. 1973. Hyphomycetes.
Deseo -509. In. They Fungi. Vols LVA. G.-C. Ainsworth,
F. K. Sparrow & A. S. Sussman (eds.). Academic Press,
toc.) New York, NY.
Kunwar (1. OK. “Manandhar,. 0. Bess seortne lair, J.B.
1986. Histopathology of soybean seeds infected with
Alternaria alternata. Phytopathology 76: in press.
Weresub, La K., & LeClair, P.M. 1971.0n Papulaspora
and bulbilliferous basidiomycetes Burgoa and
Miniamedusa. Can. J. Bot. 49:2203-22137
‘et)
fait
MYCOTAXON
VOLeCRAVLL PD. bow bao October-December 1986
NOTES DE NOMENCLATURE CONCERNANT LES HYMENOMYCETES: IV
SUR QUELQUES EPITHETES SPECIFIQUES PREOCCUPES. 3,
Régis COURTECUISSE
33, rue de Prétre
F. 59249 - AUBERS
KEY-WORDS : Hymenomycetes, Entoloma, nomenclature, type studies.
ABSTRACT : Some new cases of preoccupied epithets are studied. The following
new names are proposed: Entoloma subaerugtnosum, E.egontt, E.largenttt, E.fla-
votdes, E.gugmantt.Some new combinations are added: E.vtrescens (Berk. and
Curt.), E.largentit var. badtodorsum (Largent), E.largenttt var. alntcola
(Largent), E.largentitvar. fartnaceum (Largent).
RESUME : Quelques nouveaux cas d'épithétes préoccupés sont étudiés. Les nou-
veaux noms suivants sont proposés: Entoloma subaerugtnosum, FE. egontt,FE.lar-
genttt,E. flavotdes, E. guamantt. Quelques nouvelles combinaisons sont ajoutéés:
E.vtrescens (Berk. and Curt.), E.largenttt var. badtodorsum (Largent), E.lar-
genttt var. alntcola (Largent), E.largentttvar. fartnaceum (Largent) .
INTRODUCTION :
Cette note fait suite au travail publié précédemment (COURTECUIS-—
SE, 1984), et prend en considération quelques é€pithétes préoccupés et quelques
problémes de nomenclature dans le genre Entoloma (Fr.)Kummer. Des études de ty-
pe sont proposées pour étayer les remaniements nomenclaturaux.
REMERCIEMENTS
Nous tenons a remercier trés sincérement les directeurs des herbiers et
muséums suivants, qui nous ont aimablement accordé le prét des types et espé-
ces €étudiés: ENCB, FLAS, HSC, NY, NYS, TMI; de méme que les collégues qui, a
des titres divers nous ont permis la réalisation de ce travail: M.Bon (Saint-
Valery-sur-Somme), J.Géhu-Franck (Lille), G.Guzman (Xalapa- Mexique), J.W.
Kimbrough (Gainesville- USA), T.Kuyper (Leiden- Hollande), D.L.Largent (Arca-
ta= USA), E.Nagasawa (Tottori- Japon), M.E.Noordeloos (Leiden- Hollande), C.T.
Rogerson (Bronx- USA), J.Trimbach (Nice) et C.VanHaluwyn (Lille).
ENUMERATION ALPHABETIQUE DES EPITHETES CONSIDERES
1. AERUGINOSUM
128
an Leptonta aeruginosa Peck 1889 Bull.Torr.Bot.Cl. 26:65
Cette espéce est étudiée par HESLER (1963:327) et par LARGENT (1977:
162). Elle se place dans la section des Virtdtcaules Largent 1974, appartenant
au sous-genre Leptonta (Fr.)Noordeloos, section Paludocybe (Largent) Noordeloos
et semble bien individualisée au vu des descriptions publiées par ces deux au-
teurs.
Nous avons examiné le type de ce taxon (NYS) et nos observations corres-
pondent a celles de HESLER et de LARGENT. Nous donnons simplement un dessin
des spores de cet exemplaire type (Fig.1).
Une combinaison dans le genre Entoloma (Fr.)Kummer est impossible en rai-
son de l'existence de 1'Entoloma aeruginosum Hiroe (voir ci-dessous). Un nou-
veau nom est donc nécessaire. Nous proposons:
Entoloma_subaeruginosum Courtecuisse “om.nov.
= Leptonta aeruginosa Peck 1889 Bull.Torr.Bot.Cl. 26:65
= Leptontella aeruginosa (Peck)Murrill 1917 W.am.Fl. 10:89
non Entoloma aerugtnosum Hiroe 1939 Appl.Mushr.Set. 4:1
UD Oe
JOO
Fig.1 - Entoloma subaerugtnosum Court. (Leptonta aeruginosa Peck - Type NYS)
Spores
Cette espéce a été assez abondamment étudiée, de méme que son grou-
pe, principalement par HORAK (1971, 1973, 1978, 1980). Une synonymie trés
importante -existe, et nous la reproduisons ci-dessous, non sans en discuter
préalablement quelques points particuliers.
Le binéme le plus ancien correspondant 4 ce taxon semble étre Agartcus
vtrescens Berk. et Curt. 1860. HORAK (1976:200) a combiné ce binéme dans le
genre Entoloma en citant un basionyme incomplet et imprécis. Nous pensons u-
tile de valider ci-dessous cette combinaison. #
Nous avons examiné le type de Agartcus vtrescens Berk. et Curt. (K872).
Celui-ci est en assez mauvais état, en particulier sur le plan microscopique
et seules les spores peuvent étre observées de fagon convenable.
129
Agartcus virescens Berk. et Curt.; Type K.872; Bonin Isles; Herb.Mycol.Berk.
1879.
Spores de forme cubique, X 8,8-11 p- (Fig.2)
Gono ©
gags ®
Fig.2 - Entoloma virescens (B.et C.)Horak c.n. (Agaricus virescens B.et C. -
Type K) Spores.
Il nous semble trés regrettable que les autres particularités anatomiques
de ce champignon ne puissent é6tre observées. En effet, nous avons obtenu le
prét de deux récoltes de Entoloma aerugitnosum Hiroe, synonyme d'Ag.vtrescens
d'aprés HORAK, grace a4 l'obligeance de E. NAGASAWA (le type en aurait été per-
du d'aprés le Dr. HIROE), et quelques détails nous semblent différer des ob-
servations que nous avons faites sur Ag.vtrescens d'une part, et des descrip-
tions données par ROMAGNESI pour son Rhodophyllus holocyaneus d'autre part,
autre espéce placée en synonymie de Ag.vtrescens par HORAK.
Donnons tout d'abord une description des spécimens de E.aeruginosum Hiroe.
Entoloma aerugtnosum Hiroe; TMI 1945; On soil in mixed wood; Ochidani, Tottori
City; Sept.19, 1974; E.Nagasawa.
Exsiccatum de teinte vert foncé.
Spores X 9,5-11,5-(12) pp (t+apicule), de forme cuboide complexe, prenant
des allures d'étoile 4 6 branches selon l'orientation, avec les faces conve-
xes, séparées par des angles assez nets. (Fig.3)
Basides 4-sporiques, de grande taille, bouclées.
Sous-hyménium coralloide complexe.
Trame subparalléle.
Boucles présentes, fréquentes.
Cheilocystides nombreuses (Fig.4); Pleurocystides nombreuses, semblables
aux cheilocystides, mais plus petites. (Fig.5)
Pigment granuleux, rose-orange dans le sous-hyménium, mais verdatre dans
la trame et les cystides. Les granules présents dans la trame sont plus volu-
mineux que ceux des cystides (observations dans le Rouge Congo ammoniacal).
Rhodophyllus aeruginosus (Hiroe) Hongo; TMI 4300; on soil in mixed wood; Koko-
iS Ukehesopenl fenkeye Mytos\arkop vay wenkel-ye
Spores X 10-12-(12,5) p (tapicule), cuboides 4 contour plus ou moins ar-
rondi, de forme trés complexe, trés difficile A dessiner ! (Fig.6) .
Basides 4-sporiques, de grande taille, 50-60 x 10-15 p(+ stérigmates) ,
bouclées.
Sous-hyménium plus ou moins coralloide.
Trame subparalléle.
Boucles fréquentes.
Cheilocystides (Fig.7) et pleurocystides (Fig.8) présentes, 45-60 x 3,5-
10 p.
130
Fig. 3-5 - Entoloma aerugtnosum Hiroe (TMI 1945); 3: Spores; 4: Cheilocystides
5: Pleurocystides et basidioles.
Hyphes remarquablement colorées par un pigment intracellulaire granuleux
vert-jaune vif (pouvant évoquer des algues !), plus ou moins brunatre vers le
sous-hyménium. Cellules faciales a pigment identique, mais plus brun.
Il nous a semblé frappant que les spores de Agaricus vtrescens Berk. et
Curt. soient trés nettement cubiques et ne posent aucun probléme au dessin,
alors que celles de Entoloma aerugtnosum Hiroe sont de forme trés complexe,
bien que baties sur le modéle cubique, et causent de véritables difficultés
au dessinateur.
C'est pour cette raison en particulier qu'il serait trés intéressant de
pouvoir controler les autres caractéres microscopiques d'Agartcus vtrescens.
D'autre part, HORAK (1980) synonymise également Rhodophyllus holocya-
neusS Romagnesi 1941 a A.vtrescens (donc a E.aerugtnosum). Or, ROMAGNESI, dans
sa description princeps, insiste sur l'existence "dans le sous-hyménium...
d'une couche trés distincte d'hyphes gréles brun-noiratre dans 1'ammoniaque
et vert-noir dans l'acide sulphurique, sans corpuscules trés différenciés,
tranchant nettement sur le médiostrate."
Cette couche nette n'a pas pu 6tre retrouvée sur les exemplaires japo-
nais de E.aerugtnosum, provenant d'une localité citée par Hiroe dans sa pu-
blication originale et dont l'identité ne peut 6tre mise en doute. Il est
vrai que sur ces spécimens, la coloration verte de la trame passe au brun-
vert et au brun vers le sous-hyménium, mais ce passage:se fait d'une fagon
assez progressive et ne forme pas de séparation bien tranchée.
Il semble donc que la correspondance entre Agartcus vtrescens et Entolo-
ma aeruginosum d'une part, et ce dernier et Rhodophyllus holocyaneus d'au-
5:
Fig.6-8 - Entoloma aerugtnosum Hiroe (TMI 4300); 6: Spores; 7:Cheilocystides,
Sous-hyménium et tramé; 8: Pleurocystides.
tre part, ne soit pas absolument parfaite. Il est néanmoins possible que
les spores 4gées du type de Berkeley de Agartcus virescens soient déformées
par rapport a la forme plus complexe qu'elles auraient pu présenter, et aus-
si que la répartition et l'aspect des pigments puissent 6tre modifiés par
une conservation en exsiccatum (l.aerugtnosum) ou en solution formolique
(R. holocyaneus) .
Il nous semblerait utile de reconsidérer la synonymie ci-dessous, sur
la base de comparaisons d'exemplaires frais et en bon état.
Néanmoins, dans 1'état actuel des données concernant ces taxons, on peut
admettre le point de vue de HORAK et donner la synonymie suivante
Entoloma_ vtrescens (Berk. et Curt.) Horak comb.nov.
Basionyme = Agartcus virescens Berk. et Curt. 1860 Proc.Amer.
Acad.Arts Set. 4:116
U52
Entoloma virescens (Berk. et Curt.)Horak 1976 Sydowta 28:
200 (basionyme incomplet et imprécis)
? Entoloma aerugtnosum Hiroe 1939 Appl.Mushr.Set. 4:1
? Rhodophyllus aerugitnosus (Hiroe) Hongo
? Rhodophyllus holocyaneus Romagnesi 1941 Prodr.Fl.Mycol.Ma-
dagascar 2:150
Les autres synonymes donnés par HORAK sont : (pas d'échantillons
examinés) = Cortinartus hochstettert Reichart 1866 Ver. Zool. Bot.Ges.
Wren 16:376
Hygrophorus hochstettert (Reichart)Reichart 1870 Retse Fr.
Novarra :144
Entoloma hochstettert (Reichart)Stevenson 1962 Kew Bull.
Ge Asis
Hygrophorus cyaneus Berk. ap.Hooker 1867 Handb.N.Zeal.Fl.
2604
Agartcus tneanus var. polychrous Berk. et Br. 1871 J.Limn.
Soc.Bot. 11:540
i
Signalons que l'on peut trouver une icone de Rhodophyllus aerugtnosus
(Hiroe)Hongo dans IMAZEKI et HONGO (1957:77).
2. ALBIDUM
a. Entoloma_albtdum Murrill 1917 N.am.Fl. 10:119
= Entoloma ctnerascens Hesler 1967 Beth.Nov.Hedw. 23:177
= Entoloma canum Hesler 1974 Mycologta 66:717
Littérature: HESLER: 1963:318 (type study), 1967:177, 1974:717.
Cette espéce semble bien individualisée.
En 1967, HESLER propose un nouveau nom, E£.ctnerascens, pour éviter la
confusion avec Leptontella albtda Murrill 1917, qu'il venait de combiner
dans le méme ouvrage, quelques pages plus té6t, dans le genre Entoloma. C'est
en fait cette combinaison qui est illégitime (voir ci-dessous), et le nouveau
nom proposé est superflu puisque 1'Entoloma albidum Murrill est le premier a
avoir €été proposé dans ce genre, et a donc-la priorité.
De plus, en 1974, HESLER remplace son E.ctnerascens par E.canum, pour
éviter 4 nouveau une confusion hypothétique avec Leptonta ctnerascens Velenov-
ski 1921, bindéme qui, a notre connaissance, n'a jamais été combiné dans le
genre Entoloma. E.canum est donc un nomen superfluum qui en remplace un autre
non Leptonta albtda (Murrill)Murrill 1917 (voir ci-dessous)
D'aprés HORAK (1980:313), cette espéce n'appartient pas au genre Entoloma.
Littérature: HESLER (1963:328, type study, et 1967:27); LARGENT (1974:58
type study et 1977:156).
Nous avons rééxaminé le type de cette espéce:
Leptontella albtda Murrill; Type NY 1759; on soil in deep shade. Chapell Hill.
September 12, 1915; Coll. H.R.Totten.
Fiche de description manuscrite accompagnant le type: "Cap hemispheritic
to convex expanded, up to 2-3 cm wide, slightly umbilicate or depressed in
center with a little rounded umbo; margin typically not striate but sometimes
with long distant ridges. Surface finely squamulose, dull or shining, color a
very light cream tan at maturity, whitish when young. Flesh thin, soft, brit-
tle, whitish, less than 1 mm thick at stem. Taste woody and mildly bitter.
Gills not crowded, more or less sinuate, sometimes slightly decurrent by a
little tooth, about 5 mm wide near the stem, margin rather regular for the
genus, a clear flesh color. Stem 3-6,5 cm long, 1,5-2 mm thick about color
EGS
of cap all over, smooth, finely granular above, tough, distinctly hollow.
Spores salmon flesh color, broadly ellip.irreg., decidedly angular apiculate.
rose-coloured: 8-9 X 6 p.
Nos observations sont les suivantes
Spores. 9=11,5 X 6,5-7,6 2 (+ apicule), A angles nets et saillants, 4a
contour assez courtement elliptique, 5-6@-gones (Fig.9).
Sous-hyménium pavimenteux a petites cellules aplaties en 5 a 6 couches.
Trame subparalléle.
Laticiféres présents (rares).
Boucles nulles.
Cuticule avec un épicutis présentant des fascicules d'hyphes un peu bou-
dinées a extrémité cylindro-clavée , a4 pigment vacuolaire (Fig.10); laticifé-
res présents (rares).
Nos mesures de spores sont un peu plus grandes que celles de HESLER et de
LARGENT.
: Ce taxon, qui semble bien individualisé, entre dans le sous-genre Lepto-
nta (Fr.)Noordeloos, section Paludocybe (Largent)Noordeloos. Une révision de
ce groupe est toujours absente de la littérature, et on ne peut que citer la
position que LARGENT‘attribue a cette espéce, dans la section Albtdicaules
Largent 1974 (Paludocybe étant alors au rang de sous-genre).
Un nouveau nom a déja été proposé pour cette espéce par Morgan-Jones
Entoloma_heslert Morgan-Jones 1971 Can.d.Bot. 49:1052
= Leptontella albtda Murrill 1917 N.am.Fl> 10:87
= Leptonta albtda (Murrill)Murrill 1917 Mycologta 9:180
= Entoloma albtdum (Murrill)Hesler 1967 Beth.Nov.Hedw. 23:27
non Entoloma albtdum Murrill 1917 W.am.FLl. 10:119 -
Cependant, le nom Entoloma heslert fut a nouveau employé 5 ans plus tard
par Horak pour désigner une autre espéce de Hesler, au nom lui aussi préoccu-
pé: Entoloma gractle Hesler. Cette espéce, qui appartient a la section Stau-
rosport telle qu'elle est décrite par Romagnesi en 1941 et posséde donc des
spores "cuboides" n'a rien 4 voir avec E.heslert Morgan-Jones, placé comme
nous venons de le voir dans la section Paludocybe du sous-genre Leptonta.
Nous proposons pour cette espéce le nouveau nom suivant :
Entoloma egontt_Courtecuisse “om.nov.
= Entoloma gractle Hesler 1967 Beth.Nov.Hedw. 23:25
non Entoloma gractle Stevenson 1962 Kew Bull. 16:236
= Entoloma heslert Horak 1976 Sydowta 28:216
non Entoloma heslert Morgan-Jones 1971 Can.J.Bot. 49:1052
3. CONVEXUM
a. Entoloma_convexum_Stevenson 1962 Kew Bull. 16:235
Ce taxon est une bonne espéce.
Références: HORAK (1971:425, 1973:28, 1980:136).
Cette espéce est créée en 1977 par LARGENT, ainsi que trois variétés.
Elle représente une bonne espéce, et un nouveau nom doit é6étre proposé puis-
que son introduction dans le genre Entoloma est rendue impossible par 1'ho-
monyme antérieur sus-cité. Les types de cette espéce et de ses variétés nous
ont révélé les caractéres suivants:
Leptonta convexa Largent; Type (HSC) DLL 7021; gregarious on needle humus on
fallen branches of sitka spruce. Big Lagoon Forest. Humbolt Co., Ca.; 9 Déc.
1974; Coll. and Det.: Largent.
Fig. 9-10 - Entoloma heslert Morgan-Jones (Leptontella albtda Murrill - Type
NY); 9: Spores; 10:Cuticule.
Spores 8,5-10 X 5,5-6,5 p, de silhouette elliptique, de ee Comp less
A nombreuses facettes plus ou moins bien marquées, avec une dépression supra-
apiculaire. Apicule petit. (Fig.11)
Basides 4-sporiques, bouclées, clavées, fines, é6lancées. (Fig.12)
Boucles présentes, mais assez rares dans la trame.
Sous-hyménium celluleux régulier, a cellules plus ou moins cubiques.
Trame paralléle, a cellules relativement courtes, jusque 45 X 6 p, plus
fines juste sous le sous-hyménium.
Laticiféres non observés dans les lames.
Cheilocystides et pleurocystides nulles. Marge garnie de basidioles.
Trame piléique congophobe, a hyphes moyennes ou fines. Epicutis formé de
chaines de cellules cylindracées, jusque 35-60 X 10-21 Pp, en fascicules un peu
redressées. Pigment vacuolaire. Segment terminal des chaines d'hyphes non dif-
férencié.
Laticiféres assez rares dans la cuticule. (Fig.13)
Nous proposons pour cette espéce le nouveau nom suivant :
Entoloma largenttt Courtecuisse nom.nov.
= Leptonta convexa Largent 1977 Btbl.Mycol. 55:11
non Entoloma convexum Stevenson 1962 Kew Bull. 162.235
Leptonta convexa var. badtodorsa Largent; Holotype (HSC) L 3253; scattered in
mossy humus beneath Vine mapple and near Douglas fir; Tenino Mounts; Piearce
Co., Wash.; 28 Oct.1967.
135
Fig.11-13 - Entoloma largenttt Court. (Leptonta convexa Largent - Type HSC)
Mimispores; ales baside rhs: CuLLcuLe
Spores 8,5-11 X 5,5-6,5-(6,7) p, de silhouette ellipsoidale, de forme as-
sez complexe, a angles nombreux et peu marqués. Apicule petit. (Pig.14)
Basides 4-sporiques, bouclées, cylindracées plus ou moins clavées, moyen-
nement trapues (Fig.15)
Boucles présentes dans l'hyménium et le sous-hyménium, nulles ou trés ra-
res dans la trame.
Sous-hyménium mince, confus, subcelluleux a pavimenteux.
Trame paralléle a subparalléle, formée d'hyphes moyennes.
Cuticule: Epicutis formé de chainettes en boudin ou en chapelet de saucis-
ses, en fascicules, a pigment vacuolaire abondant. Eléments souvent courts et
trapus, par exemple 30-70 X 10-30 p (Fig.16)
Laticiféres non observés.
La combinaison suivante est proposée
Entoloma largenttt var. badtodorsum (Largent)comb.nov.
Basionyme = Leptonta convexa var. badtodorsa Largent 1977 Brbl.
Mycol. 55:114
Le rang variétal semble pouvoir 6tre conservé pour le moment, car plusieurs
caractéres macroscopiques et é6écologiques différent par rapport au type (cf.
LARGENT 1.c.). Sur le plan microscopique, les spores semblent un peu plus lon-
gues et les éléments de 1'épicutis un peu plus larges.
136
Fig.14-16 - Entoloma largentit var. badtodorsum (Largent)Court. (Type HSC)
14: spores, 15:basides, 16: éléments de 1'épicutis.
Leptonta convexa var. alntcola Largent; Type (HSC) L 1486; gregarious in a hu-
mus of alder leaves; Friday Harbor, San Juan Co., Wash.; 14 Nov. 1965; Coll.
and Det.: Largent.
Spores 8-10,5 X 5,5-6,5 p, de forme elliptique allongée, obscurément angu-
leuses, de forme complexe, semblant en moyenne les plus étirées du groupe (mé-
me si les fourchettes de dimensions sont comparables a celles des autres taxons,
le rapport L/1l moyen semble plus elevé) (Fig.17)
Basides 4-sporiques, bouclées (Fig.18)
Boucles trés rares, sauf au pied des basides.
Sous-hyménium celluleux pavimenteux serré.
Trame subparalléle. Hyphes moyennes, plus grosses dans la médiostrate.
Cheilocystides et pleurocystides nulles. Laticiféres non observés.
Cuticule: Epicutis avec des chainettes d'hyphes boudinées 4 éléments sou-
vent trés larges, 40-90 X 15-40 p, en fascicules plus ou moins dressés. Pigment
vacuolaire, parfois presque invisible, semblant plus pale que chez les autres
membres du groupe. (Fig.19)
La combinaison suivante est proposée :
Entoloma largentit var. alnteola (Largent)comb.nov.
Basionyme = Leptonta convexa var. alntcola Largent 1977 B¢bl.
Mycol. 55:115
E37
Fig.17-19 - Entoloma largenttt var. alnicola (Largent)Court. (Type HSC)
17: spores, 18: baside, 19: éléments de la cuticule.
Fig.20-21 - Entoloma largentit var. farinacea (Largent)Court. (Type HSC)
20: spores, 21: basides.
138
Le rang variétal est ici encore conservé, bien que les différences sem-
blent plus importantes par rapport au type que pour la var. badtodorsum. u'
habitat est caducicole, les spores apparemment plus étirées et plus obscuré-
ment anguleuses, les éléments de 1'épicutis beaucoup mains pigmentés et plus
larges. D'autres récoltes pourraient permettre une élévation au rang spécifi-
que si ces caractéres se trouvaient constamment associés.
Leptonta convexa var. fartnacea Largent; Type (HSC) DL 3148; solitary in mas-
ses near conifers; Ipsut Campground. Mt.Rainier Nat.Park, Piearce Co., Wash.
14705-21965.
Spores 8-9,5 X 5,5-6-(6,5) p, de forme ellipsoide, a4 angulation assez
nette, 5-7-gones, assez proches de celle du type, mais peut-&étre un peu plus
confuses. Apicule petit. (Fig.20)
Basides 4-sporiques, cylindracées, un peu clavées, assez courtes et tra-
pues, bouclées. (Fig. 21)
Boucles trés rares ou nulles dans la trame.
Sous-hyménium celluleux a petits éléments serrés.
Trame paralléle a hyphes moyennes.
Cheilocystides et pleurocystides nulles. Laticiféres nuls.
Cuticule non observée (exemplaire trop fragmenté).
La combinaison suivante est proposée :
Entoloma largenttit var. farinaceum (Largent)comb.nov.
Basionyme = Leptonta convexa var. fartnacea Largent 1977 Brbl.
Mycol. 55:117
Le rang variétal est conservé, bien qu'il puisse ne s'agir que d'une for-
me de Entoloma largenttt.En effet, les spores semblent les plus proches de
celles du type. Sur le plan écologique, les coniféres forment le dénominateur
commun. Néanmoins, nous n'avons pu observer la cuticule de ce taxon, et il
semble que les couleurs et l'aspect macroscopique soient quelque peu diffé-
rents de ceux du type. D'autres récoltes permettront d'apporter un élément de
réponse a ce probléme.
4. CYANEUM
a. Agartcus ; cyaneus Peck 1873 Bull. Buff.Soe.Nat.Set. 1:49
non Agartcus cyanéus Bull. 1783 Herb.Fr. n°170 (= Stropharta)
= Entoloma eyaneum Saccardo 1887 Syll.Fung. 5:692 (ut E.cyaneum Peck)
= Leptonta cyanea ("Peck")Mazzer ap.Largent 1977 Bitbl.Mycol. 55:66
Sur le plan purement nomenclatural, le basionyme de Peck (1873) étant
préoccupé par l'Agartcus cyaneus de Bulliard (1783), c'est Saccardo qui endos-
se la paternité de l'espéce, ayant publié la premiére combinaison valide.(Art.72.1)
Sur le plan taxonomique, la situation est beaucoup plus confuse. En effet,
HESLER (1963:351) synonymise cette espéce avec Entoloma vtolaceum Murrill 1917,
mais LARGENT (1977:122) écrit que cette derniére espéce est un mtxtum compost-
tum formé de 1'E.cyaneum Sacc. et de E.porphyrophaeum (Fr.)Karsten. D'ailleurs
HESLER (1963:351) ne note pas de boucles chez E.vtolaceum, alors que te type de
A.cyaneus Peck en est pourvu (voir plus loin). Nous admettons que £.vtolaceum
doit 6tre abandonné en tant que nomen confusum.
L'examen du type de Agartcus cyaneus nous a revelé les caractéres suivants:
Agartcus cyaneus Peck; Type (NYS); Shandaken-Worcester; Leg. C.L.Peck.
Spores petites, ovoides, 4 un peu étirées, 7,5-9,5 X 5,5-6,5 p (+ apicu-
le) . (Fig. 22)
Basides bouclées, 4-sporiques, longues, effilées, cylindroclavées.
Boucles présentes dans la trame des lames.
Sous-hyménium subcelluleux, plus ou moins coralloide a coralloide.
Trame subparalléle a paralléle, a cellules courtes et trapues dans la
médiostrate.
Pleurocystides nulles.
139
Cheilocystides i- ou 2-cellulaires, en bouquet a l'extrémité de la trame,
rendant l'aréte stérile, cylindracées 4 ventrues ou pluriétranglées, 22-45 X
5-8 p, parfois bouclées. (Fig. 23)
Sere O76
VU ang
Fig.22-23 - Entoloma cyaneum Sacc. (Agartcus cyaneus Peck - Type NYS);
22: spores, 23: cheilocystides.
La position taxonomique de cette espéce semble assez claire; une aqua-
relle figure dans la collection type, qui montre un champignon au pied rela-
tivement trapu, €paissi vers la base, de couleur bleu-violet et palissant
vers le bas ou il est lavé de bleudtre et méme de jaundtre a l'extréme base,
et squamuleux au sommet. Les lames sont d'abord blanches et le chapeau est
beige-violacé au centre, violet foncé a la marge chez le jeune imbu, et
violet opaque au disque, violet foncé a la marge chez l'adulte (sec ?).
LARGENT (1977:68) signale que MAZZER pense que ce champignon est iden-
tique a Entoloma dichroum (Pers.:Fr.)Kummer des auteurs européens et se ran-
ge a ce point de vue. Celui-ci nous semble néanmoins difficile a admettre
puisque E.dtchroum n'a pas le pied pelucheux de A.cyaneus et que ses spores
sont beaucoup plus grandes: (9)-9,3-12 X 7-9,3-(10) p selon NOORDELOOS
(1982:462). Les spores que nous avons mesurées sur le type de A.cyaneus
sont de taille comparable 4 celles observées par LARGENT (1977:67). Entoloma
eyaneum Sacc. se rangerait donc de facon trés satisfaisante dans le sous-genre
Leptonta, section Leptonta telle qu'elle est définie par NOORDELOOS (1982:
453) et viendrait 4 proximité immédiate des espéces du "complexe-dtchroum"
en tant que quatriéme membre de celui-ci (avec E.dichroum, E.allochroum
Noord. et E.tjalltngtorum Noord.) différencié par ses petites spores et aussi
par ses caractéres macroscopiques qu'il serait d'ailleurs fortement souhaita-
ble de préciser sur du matériel frais.
Il faut encore signaler que LARGENT (1977:66) cite comme synonyme supplé-
mentaire Leptontella acertcola Murrill 1917 W.am.Fl. 10:88, de méme que Ento-
Loma velatum Hesler 1967 Beth.Nov.Hedw. 23:114, malgré quelques différences
micrographiques (spores et/ou cheilocystides, etc...).
140
= = Fhodophy las cyaneus (Murrill) Singer 1947 Mycologta 39:187
= Entoloma cyaneum (Murrill)Hesler 1967 Beth.Nov.Hedw. 23:13
non Entoloma cyaneum Saccardo 1887 Syll.Fung. 5:692
Le type de Claudopus cyaneus (FLAS 17841) est constitué par une enveloppe
dans laquelle se trouvent du sable et des débris ligneux parmi lesquels nous n'
avons pas pu trouver de champignon. Nous devons donc nous référer 4 la des-
cription publiée par HESLER (1963:331 et 1967:13). D'aprés celle-ci, le cham-
pignon de Murrill ne semble pas rentrer dans le sous-genre Claudopus tel qu'il
est défini par NOORDELOOS (1981a:147), mais plut6t, peut-étre, dans le sous-
genre Paraleptonta (Rom.)ex Noord. (1.c.:149), par l'absence de pigment mem-
branaire que l'on peut imaginer puisque Hesler dit (1967:14): "cuticle repent
with colourless hyphae".
Les caractéres de cette espéce semblent par ailleurs trés originaux. La
couleur bleue en particulier est remarquable dans les entolomes 4 "port pleu-
rotoide" de l'ancien genre "morphologique" Claudopus. Cependant, HORAK (1980:
33) cite une autre espéce bleue: Claudopus cyanomelaenus Boedijn 1929 Rec.Trav.
Bot.Neerl. 24:419.
Ce dernier taxon rentre beaucoup plus facilement dans le sous-genre Pa-
raleptonta (Romagn.)ex Noord. Le pigment vacuolaire est cité dans la diagno-
se originale, et écarte les Claudopus ss.str.
Les deux espéces ne semblent pas différer de facgon significative.
Le stipe est trés court chez C.cyanomelaenus Boedijn: 0,5-1,5 X O,5 mn,
et peut 6tre beaucoup plus long chez C.cyaneus Murrill, jusque 3 cm X 1,5 mm
(!) d'aprés HESLER, mais nous nous demandons si ces mensurations ne peuvent
pas provenir d'une erreur de transcription, car nous avons noté, sur ]'éti-
quette présente dans le sachet du type: "3 X 1,5 mm".
L'odeur et la saveur sont données comme farineuses pour C.cyaneus, mais
aucun caractére organoleptique n'est signalé pour C.cyanomelaenus.
Les spores sont de dimensions apparemment semblables, puisque HESLER
donne’ tout @abord(1963):331)) 9=-10-(12) "xX 6-7 pu, puis” (1967214)) 8-10=(12) 7x
6-7 p (il est a noter que le sachet du type renferme une annotation de Hes-
ler, portant: sp. 9-12,5 X 6-7,5 p !), ceci pour C.cyaneus et, que HORAK
(1980:33) cite 9-11 xX 5,5-6,5 p pour C.cyanomelaenum qui sont les mesures
données dans la diagnose originale. De plus, la forme de la spore photogra-
phiée par HESLER (1963:356, fig.19) correspond de fagon satisfaisante a la
fig.9 de Boedijn (1929:418).
Néanmoins, comme nous n'avons pas pu examiner de matériel de ces deux
espéces, il semble prématuré de les proposer comme synonymes. Si 1'examen
des carpophores frais pouvait révéler cette identité, Claudopus cyanomelae-
nus Boedijn 1929 devrait &tre combiné dans Entoloma. Dans le cas contraire,
il faudrait proposer un nouveau nom pour le Claudopus cyaneus Murrill.
S- FLAVIDUM
as Clitoptlus_ flavidus Massee 1908 Kew Bull.:4
= Entoloma flavidum (Massee)Corner et Horak ap.Horak 1980 Beth.Nov.
Hedw. 65:108
Ce taxon, décrit et figuré par HORAK (1980:108) semble assez polymorphe
si l'on admet que le type et les exemplaires récoltés a4 Singapour par CORNER
sont conspécifiques.
Le type de cette espéce nous a révélé les caractéres suivants :
Clttoptlus flavidus Massee; Type (K); Singapour; Ridley 56D (exemplaires trés
fragmentés !)
Spores 8,5-10 X 6,5-7 p (+ apicule), assez petites, de forme simple et
a angles moyennement marqués (Fig.24).
Basides 4-sporiques, semblant non bouclées (?). L'hyménium s'est montré
étre en trés mauvais état, confus, la majorité des cellules étant collapsées.
Boucles non observées avec certitude.
141
Sous-hyménium semblant plus ou moins coralloide.
Trame subparalléle a hyphes moyennes a petites.
Aréte des lames et cuticule non observées en raison de l'état de fraction-
nement de l'exemplaire.
Les mesures sporales correspondent a celles données par HORAK, mais sont
trés différentes de celles de la description originale que nous reproduisons
ici :
"Ptleus carnosus, e convexo umbtltcatus, vel tnfundtbultformts, margine
undulatus vel lobatus, laevis, minute squanulosus, flavtdus, 4-5 cm latus.
Lamellae subeonfertae plus minusve, decurrentes, albotnecarnatus, Sporae elltp-
tteae, laxe et minute asperulae, roseotinctae, 6-7 X 4-4,5 yp. Bastdta clavata
21-25 X 6-7 yp. Stipes farctus, subftbrillosus bast tnerassatus lacunosus, ptleo
concolor, 3-4 em longus, 1 em crassus".
Po OW
fo) ©
Fig.24 - Entoloma flavidum (Massee)Corner et Horak (Clitoptlus flavidus Massee.
Type K), spores.
Le type de cette espéce nous a révélé les caractéres suivants :
Eectlta flavtda Peck; Type (NY); Stow.,Mass.; Coll. G.E.Morriss; 11.08.1908
Spores 7,5-9,2 X 5,5-6 p (+ apicule), allongées, ovoides, nettement angu-
leuses avec une dépression supra-apiculaire souvent nette (Fig.25).
Basides 4-sporiques, non bouclées.
Boucles non observées.
Sous-hyménium subcelluleux, plus ou moins coralloide a petites cellules
serrées.
Trame subparalléle a plus ou moins emmélée irréguliére.
Pleurocystides nulles.
Cheilocystides rares apparemment, plus ou moins collapsées, de forme gé-
nérale cylindracée, quelques unes clavées, non mesurables (abimées) (fig.26)
Nous ne connaissons pas les caractéres macroscopiques de cette espéce,
mais elle semble s'inscrire dans le sous-genre Nolanea (Fr.)Noordeloos, section
Endochromonema (Largent et Thiers)Noordeloos. Les sous-sections Icterina
Noordeloos et Chetlocysttdtata Noordeloos pourraient accueillir ce taxon. Au-
cune espéce ne semble lui correspondre.
Nous proposons le nouveau nom suivant
Entoloma flavotdes Courtecuisse ”om.nov.
= Ecetlta flavtda Peck 1909 Bull.Torr.Bot.Cl. 36:153
non Entoloma flavtdum (Massee)Corner et Horak ap. Horak 1980
Beth. Nov.Hedw. 65:108
142
2 ie
S07 —
Fig.25-26 - Entoloma flavotdes Court. (Eectlta flavida Peck - Type Ny)
25: spores; 26: cheilocystides.
6. MEXICANUM
a. Ne ACN aA ANS 1917 N.am. Fl. 10:85
= Entoloma subcaeruleum Hesler 1967 Beth.Nov.Hedw. 23:142
= Leptonta subeaerulea (Hesler)Largent 1977 Brbl.Mycol. 55:132
voir aussi HESLER (1963:339), type study.
_—— i ———$ ee
= Entoloma mextcanum (Murrill)Hesler 1967 Beth.Nov. Hedw. 23795
Nous n'avons pas vu le type de cette espéce, 6étudié par HESLER (1963:339)
puis LARGENT (1977:259) que ce dernier synonymise a4 Leptonta undulatelka
(Peck) Saccardo. Cependant, les spores telles qu'elles ont été mesurées par
HESLER et par LARGENT nous semblent un peu petites par rapport a celles que
nous avons observées sur le type de Agartcus undulatellus Peck : (7,5) -8-
10 X 6-7 p. Nous préférons rester, pour l'instant, prudent quant a cette sy-
nonymie.
c. Rhodophyllus mextcanus_Guzman 1975 Bol.Soc.Mex.Mte. 9:63
Le type de cette espéce nous a montré les caractéres suivants
Rhodophyllus mextcanus Guzman; Tipo (ENCB); Carretera Montemorelos 4 Linares
cerca de Arroyo de Encadenados, Nuevo Leon. Agosto 7, 1973; En suelo solita-
rio matorral microfilo con Telochtstes chrysophtalmus (L.)Th.Fr.; Coll. et
Det.: G.Guzman, n°11216.
Spores subsphériques a trés légérement ovoides, a angles et facettes
nettes, souvent 6-gones, 9,5-10 X 8,5-9,5 p (+ apicule) (Fig.27).
Basides 4-sporiques, non bouclées. Une sclérobaside observée, un peu
plus petite.
143
Sous-hyménium celluleux, coralloide a petits éléments noduleux.
Trame subparalléle a hyphes fines. Boucles nulles.
Pleurocystides non observées (les minuscules cystides faciales signa-
lées dans la diagnose latine par Guzman semblent difficiles a voir, ou peut-
étre diminuer encore de taille sur exsiccata ?)
Cheilocystides plus ou moins lagéniformes, ventrues, plus ou moins mu-
cronées. (Fig. 28)
Cuticule a 6picutis plus ou moins hyméniforme, les hyphes terminales étant
redressées, cylirndro-clavées et obtuses (Fig.29). Pas de boucles.
OVORGO®
28
29
Fig. 27-29 - Entoloma guzmantt Court. (Rhodophyllus mextcanus Guzman - Type
ENCB); 27: spores, 28: cheilocystide-; 29: cuticule.
La position taxonomique au sein du genre Entoloma (Fr.)Kummer nous semble
trés originale. Le sous-genre Allocybe Noordeloos pourrait convenir d'aprés
la clé présentée par cet auteur (1981b:135), mais la diagnose du sous-genre
ne permet pas de confirmer cette appartenance; les boucles sont présentes au
pied des basides et la cuticule est un cutis banal. Deux caractéres trés im-
portants qui séparent le Rh.mextcanus Guzman de ce sous-genre.
Aucun taxon supra-spécifique ne semble pouvoir accueillir cette espéce
remarquable pour laquelle nous proposons le nouveau nom suivant
Entoloma guzmantt_Courtecuisse nom.nov.
= Rhodophyllus mextcanus Guzman 1975 Bol.Soc.Mex.Mic. 9:63
non Entoloma mextcanum (Murrill)Hesler 1967 Beth.Nov.Hedw.
235295
7. TENUIPES
Références: HESLER (1963:350) type study.
Nous avons examiné le type de ce champignon.
Eeetlta tenutpes Murrill; Type (NY 1466); on the ground in oak woods at
Bound Brooks (New Jersey); Coll. Earle; 6.07.1903
Une description est donnée sur une fiche manuscrite accompagnant le type:
"gregarious. Pileus 1/2 cm, convex umbilicate, tan, glabrous; mangin
144
deeply striate; lamellae decurrent, rather broad, pallid to pale pink. Spores
pink. Stipe 60 X 1 mm cylindrical glabrous concolorous. Spores obliquely api-
culate, rose coloured, 10-12 X 6-8 p."
Spores 9,5-12 X 7-8,5 p, de forme assez simple et accusée, ellipsoide,
5-6-gones, avec un apicule assez volumineux (Fig.30)
Basides 4-sporiques, non bouclées.
Sous-hyménium confus.
Trame subparalléle.
Cheilocystides et pleurocystides nulles.
Boucles nulles.
Cuticule: cutis banal Aa pigment vacuolaire, formé d'hyphes courtes.
VD OO0e
ee
Fig.30: Eectlta tenutpes Murrill - Type NY; spores.
La combinaison de caractéres: pigment non incrustant et boucles absen-
tes confére a cette espéce une position intéressante et assez difficile 4a
interpréter. La place la plus vraisemblable que l'on puisse lui attribuer
semble se situer au sein du sous-genre Omphalopsts Noordeloos (= Eectlta
(Fr.)Q. ss.Romagnesi 1974). Seul Entoloma omphaltaeformis (Vel.)Noordeloos
semble dépourvu de boucles dans ce sous-genre. La description de Velenovsky
(1947:79) et l'étude microscopique de NOORDELOOS (1979:261) ne semblent d'
ailleurs nullement en contradiction flagrante avec les renseignements recueil-
lis au sujet du champignon américain de Murrill. Des études complémentaires
semblent nécessaires avant de proposer une synonymie entre ces deux taxons
ou un nouveau nom pour l'espéce du nouveau continent. Une combinaison dans
le genre Entoloma serait en effet impossible en raison de l'existence d'un
homonyme antérieur (voir ci-dessous).
Références: HESLER (1963:350)
Cette espéce est de position inconnue, autant que nous le sachions.
c. Nolanea tenuipes orton 1960 Trans.Brit.Mycol.Soc. 43:334
Le nouveau nom Entoloma Leptopus Noordeloos 1980 Persoonta 10(4):442 a
été proposé pour cette espéce.
Nous menons actuellement une étude comparative entre E.leptopus et E.
hebes (Romagnesi)Trimbach dont les résultats seront publiés ultérieurement.
BIBLIOGRAPHIE :
Berkeley M.T. et M.A. Curtis - 1860 - Characters of new fungi collected in the
North Pacific exploring expedition by Ch.Wright. Proc. Amer.Acad.Arts Sct.
4:111-130
145
Boedijn K.B. - 1929 - Beitrag zur Kenntniss der Pilzflora von Sumatra. Rec.
Trav.Bot.Neerl. 26:396-433
Courtecuisse R. - 1984 - Notes de nomenclature concernant les Hyménomycétes:
sur quelques épithétes spécifiques préoccupés. I. Doc.Mycol. 54-55:73-
92 (Lille - St.Valery sur Somme)
Guzman G. - 1975 - Un nuevo genero y dos nuevas especies de agaricaceos mexi-
canos. Bol.Soc.Mex.Mic. 9:61-66 (Mexico)
Hesler L.R. - 1963 - A study of Rhodophyllus types. Brittonta 15 (4) :324-366
Hesler L.R. - 1967 - Entoloma in southeastern North America. Beth.WNov.Hedw.
23:196 pp. + 240 figs.
Hesler L.R. - 1974 - Name corrections in Entoloma. Mycologta 66:715-717
Hiroe I. - 1939 - Entoloma of Japan (2). Notes on larger fungi on the San-in
District (VII). Appl.Mushr.Set. 4(1):1-3
Horak E. - 1971 - A contribution towards the revision of the Agartcales (Fun-
gi) from New Zealand. N.Zeal.d.Bot. 9:403-462
Horak E. - 1973 - Fungt Agartcint Neozelandiae-.. 1. Entoloma (Fr.) and related
genera. Beth.Nov.Hedw. 43:1-86
Horak E. - 1978 - On cuboid-spored species of Entoloma (Agartcales). Sydowta
28:171-236 (Horn)
Horak E. - 1980 - Entoloma in Indomalaya and Australasia. Beth.Nov.Hedw. 65:
S5ZT0p.
Imazeki R. et T. Hongo - 1957 - Coloured Illustrations of Fungi of Japan.
Nolet cant Sl pp.
Largent D.L. - 1974 - Rhodophylloid fungi from the pacific coast (United Sta-
tes) IV. Infrageneric concepts in Entoloma, Nolanea and Leptonta.
Mycologta 66 (6) 987-1021
Largent D.L - 1974 - Studies in the rhodophylloid fungi. V. Leptonta subgenus
Paludocybe section Albtdtcaules and section Rosetcaules and related
taxa. Northw.Set. 48(1) :57-65
Largent D.L. - 1977 - The genus Leptonta on the pacific coast of the United
States. Btbl.Mycol. 55:286 pp. + 94 figs.
Massee C. - 1908 - Kew Bull. :4
Noordeloos M.E. - 1979 - Type study in entolomatoid species in the Velenovsky
herbarium.I. Persoonta 10(2):245-265 (Leiden)
Noordeloos M.E. - 1980 - Entoloma subgenus Wolanea in the Netherlands and
adjacent regions with a reconnaissance of its remaining taxa in Europe.
Persoonta 10(4):427-534 (Leiden)
Noordeloos M.E. - 1981a- Introduction to the taxonomy of the genus Entoloma
sensu lato (Agaricales). Persoonta 11(2):121-151 (Leiden)
Noordeloos M.E. - 1981b- Entoloma subgenus Entoloma and Allocybe in the Nether-
lands and adjacent regions with a reconnaissance of their remaining taxa
in Europe. Persoonta 11(2):153-256 (Leiden)
Noordeloos M.E. - 1982 - Entoloma subgenus Leptonta in Northwestern Europe. I.
Introduction and a revision on its section Leptonta. Persoonta 11(4) :451-
471 ( Leiden)
Romagnesi H. - 1941 - Les rhodoohylles de Madagascar, Entoloma, Nolanea, Lep-
tonta, Ecctlta, Claudopus, avec une introduction générale sur la classi-
fication, la phylogénie, la répartition géographique et la toxicité des
Rhodogontosporales. Prodr.F1l.Mycol.Madag. 2:164 pp.
Velenovsky J. - 1947 - Wovttates Mycologtcae Novtsstmae. Op.Bot.Cech. 4:167 pp.
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MYCOTAXON
ViGte xh. ip ee p53 October-December 1986
TWO NEW DEMATIACEOUS HYPHOMYCETES
FROM TROPICAL FOREST LITTER
Ibpenbugey, auvereveroial, Ylaveh Sai rkhwceirove) Ohevenzioul
Laboratorio di Micologia, Dipartimento di Biologia Vegetale’,
Universita di Roma, Largo Cristina di Svezia 24, 00165 Roma,
Italy
Abstract
Descriptions and_illustrations of two new species
Ctavematiacecous Hyphomycetes, /Cirncinotrichun Fla —
gelliforme and Pyricularia sphaerulata, discovered
OM vropicalwhorestrilitrver, in«<Lvory (Coast), are siven:
Two new species of the Dematiaceous Hyphomycete genera
Circinotrichum Nees (1817) and Pyricularia Sacc. (1880) were
found onetorest ditter an) the TaijNational iPark ((ivory? Coast).
Circinotrichum flagelliforme sp.nov.
Coloniaesnyoophy lilac, punctiformes, circulares’, setis, irregu-—
Jariter dispositis circumdatae, nonnumquam coalescentes, .co-
nidiis albae. Mycelium et stroma partim superficialia partim
immersa. Setae simplices, erectae, rectae vel modice flexuo-
sae, leves, usque ad 15-septatae, in parte basSilari .crassitu
nicatae, brunneae, gradatim angustiores et rarius septatae
ad apicem rotundatum, tenuitunicatum, subhyalinum et subtilem;
usque ad 900 wm longae et 3.5 wm crassae ad basim (in basi
bulbosa 9-14.5 wm crassae). Cellulae conidiogenae polyblasti-
cae, solitariae, percurrenter proliferantes, lageniformes,
apice inflato, tenuitunicatae, subhyalinae, parallelae et ag-
gregatae, .e Sstromate subzili superficiali orientes, 7-10 x
2.3-3.5 wm. Conidia hyalina, omnia agglutinata, fusiformia
vel subfalcata, basi obtusa et apice tenui praedita, 16-20 x
2.3-2.7 pm.
inyvlOorVites senor tuus arvponitenlaut to1iae. rel, ini'Oraveboris,
16. TIT 1983), nolortypus’:GROHBYI29A; -in Tolais emortuis ‘ar
‘Doris Latitoliae, Tal, mm V7O0reveboris,--Loske.,.1983, "para
typus: ROHB 129A(1)
Colonies hypophyllous, punctiform, roundish, surrounded
by irregularly distributed setae, sometimes coalescing to
149
form irregular patches, whitish due to sporulation. Mycelium
and stroma partly superficial, partly immersed. Setae simple,
erect, straight or slightly flexuous, smooth, up to 15-septa-—
te, thicker-walled near the base, brown, progressively paler,
gradually tapering and less closely septate towards the thin-
walled, subhyaline, slender and rounded apex; up to 900 pm
long and 3.5 wm wide near the base (9-14.5 wm wide at the bul
bous base). Conidiogenous cells solitary, lageniform, swollen
at the apex, thin-walled, subhyaline, forming a dense hyme-
HNiMMmeOMelvie. thin and Superficial stroma, “7-LO xX 2.3-3.0 mM.
Conidiogenesis polyblastic with percurrent proliferation. Co-
nidia hyaline, aggregated in slimy masses, fusiform to subfal
cate, obtuse at the base and narrowing at the apex, 16-20 x
2.3=—2./7 pm.
The morphology of the conidiogenous cells and conidia
and, Ghe=presence, of unbranched setae are. the) main -characteri-
stics) that allow us to include this species in the genus Cir-
cinotrichum
Among the different, species deseribed ‘in, this: genus. only
three, C. papakurae Hughes & Pirozynski (1971), C. rigidum
Sutton (1980) and C. britannicum P.M. Kirk (1981) possess
exclusively straight sterile setae. Our species differs from
C. papakurae in having much longer and more closely septate
setae and conidia different in shape and dimensions. It dif-
fers from C. rigidum in general morphology and dimensions,
moreover “this (hast species 2s,-described :as' phialidic- rand with
conidia bearing at each end a minute appendage. C. flagel-
ivforme also differs fromuc. britannicum in having much: lLon—
ger -seétae and conidiogenous cells and, conidia different in
shape and dimensions.
For these reasons we propose C.flagelliforme as new
Species.
Nowadays many different species belong to this genus,
butthe way an which conidia are formed is a: matter of, discus
Sion. s ne only experimental data von conidial sontogceny in gthe
Circinotrichum-Gyrothrix-Ceratocladium complex have been pu-
blished by Cunningham (1974) who describes the conidiogenesis
of Ge pediculata, studied in pure culture by time-lapse .qabser
vations, as "phialidic sensu stricto". There is no evidence
of the presence of phialidic ontogeny in the previously de-
scribed species of these genera.
Later, Sutton (1980); described. the phialidic .c., rigidum,
white, also accepting the species C. fertile Pirozynski.-&
Hodges (1973) in the genus. The latter is the only species
which presents a conspicuous sympodial proliferation.
We-don"t Know 4f the conidiocenesis: in this’ complex is
phiialidic, holoblastic sympodial or by “successive rings of
conidia, as proposed by Pirozynski (1962); “but, in our opinion
150
it is problematic to maintain in the same genus phialidic
species, as C. rigidum, and holoblastic sympodial ones, as
C. fertile, Therefore, we think that, for the moment, is pre-
ferable to accept the combination Idriella fertilis (Piro-
zynski & Hodges) Matsushima, (1975) for Cc: fertile to mainiatn
separated the only species that is surely sympodial, which
moreover possesses conidiogenous cells and conidia typical
of Idriella Nelson & Wilhelm.
However the taxonomic rearrangement of this complex
awaits further studies on its .conidiogenesis.
Pyricularia sphaerulata sp.nov.
Coloniae effusae, amphigenae. Mycelium immersum, conidiopho-
ra macronematosa, mononematosa, simplicia, erecta, recta vel
modice flexuosa, solitaria, levia, crassitunicata, septata,
aureo-brunnea, apicem versus pallidiora et subtilius tunica-
ta, 90-132 x 3.6-4.8 pm; in basi expansa.7.2-12.6 pm lata,
septo circumscripta. Cellulae conidiogenae polyblasticae, in-
tegratae, terminales, sympodialiter proliferantes, denticu-
latae, tenuitunicatae; denticuli subcylindrici, ad apicem
truncati aperti, ad basim septati; T-1.4 x 1I-1.4 pm. Conitdia
Sicca, solitaria, acropleurogena, levia, 3-septata, fusifor-
mia, sub septo basilari aperta, ad apicem infundibuliformia,
Ssphaerula hyalina (circiter 5.5 wm diam) praedita; pallide
Olivaceo-brunnea, cellulis terminalibus pallidioribus quam
intermediis et protoplasmate granuloso repleta; 27.3-31.5 x
4-4.5 wm (sphaerula exclusa).
In tolids semortuis iorbornis Llatitoliacv Tai in Ore
Ebori sj 1S. 20.1983, holotypus:ROHE LoOA
Colonies effuse, amphigenous. Mycelium immersed. Coni-
diophores macronematous, mononematous, unbranched, erect,
straight or slightly flexuose, solitary, smooth, thick-walled,
septate, golden-brown, gradually paler and thinner-walled
towards the apex, 90-132 x 3.6-4.8 wm; 7.2-12.6 pm wide at
the swollen base. Conidiogenous cells polyblastic, integra-
ted, terminal, sympodially proliferating, denticulate, thin-
walled; denticles subcylindrical, flat’and open at’ the: apex;
septate at the base, 1-1.4 x 1-1.4m. Conidia dry, solitary,
acropleurogenous, smooth, 3-septate, fusiform, with an open-
ended base and a funnel-shaped apex, terminating at the tip
in a spherical hyaline body (about 5.5 wm diam), light oli-
vaceous-brown, with granular protoplasm and with the end
célls paler; thenvthe central ones; 27.3=31.5 x 4-455) 16m (ex-
cluding the spherical body).
The conidiophores are pigmented and well developed, the
conidiogenous cells are integrated, terminal, sympodially
1s 8 3
habit, sketch: bu. coni-
a.
- Pyricularia sphaerulata
2
rh Bis
diophores and conidia.
SZ
proliferating, polyblastic, denticulate. The conidia are shed
by the fracture of a small separating cell (rhexolytic seces-
sion) which leaves a subcylindrical open-ended denticle on
the conidiophore, and an open-ended portion at the base of
the conidiumy These characteristics allow us. Go -consider ene
inclusion of thiss species) in’ Pyriculariay Sacc,..
The fusiform conidia of the Hyphomycete hnere descrited
exceed the morphological limits of Pyricularia, stated as
"obclavato-pyriformia" by Saccardo (1880) in the original
diagnosis and confirmed by Ellis (1971) as "obpyriform, ob—
turbinate or obclavate". Since then in Pyricularia several
species have already been included with fusiform conidia,
such as P. fusispora (Matsushima) Zucconi, Onofri & Persfani
(1984), “andywithi ellipsoid conidia, such as F- @faurz Ply.
KiPk Ci9ss 4
A distinctive characteristic of this ‘speciesris che were
sence of a spherical body at the apex of the conidia. This is
less evident in old slides because of its rapid dissolution
inv bactice acia,
P. buloloensis Matsushima (1971) is the only species pu-
blished in Pyricularia which presents conidia with a similar
structure at the tip, described asw"calyptra gelatinosa 2 0ur
species differs from P. buloloensis in the shape and dimen-
sions of the conidia, that are fusiform instead of obclavate
and smaller. It also differs in the morphology of the spheri-
cal body which is located at the conidial apex and does not
surround it. Moreover, in our fungus the body seems to be con
tinuous with the open and swollen apical cell of the conidium.
For these reasons we propose P. sphaerulata as a new
species.
ACKNOWLEDGEMENTS
The Authors wish to thank the Minister of Scientific Re-
search of the Ivory Coast for the assistance he has given and
Dr. W., Gams, and/DraeG.Ss de ‘Hoog of ytherC 7B 5. 2for kKindiyvare-
viewing the manuscript.
REFERENCES
Cunningham, J. 1974. Avnew Gyrothrix in’ culture and a key. ve
species= Mycodogia 66: 122-129.
Ellis, M.B. 1971. Dematiaceous Hyphomycetes. Commonw. Mycol.
Inst., Kew, England.
Hughes; S.J. and K.A. Pirozynski. 1971. New Zealand Fungiers.
Beltrantella, *Circinotrichum, and Gorothrax*. ON. (aonbin
Bot. 9: 39-45.
Kirk,P.M. 1981. New or interesting microfungi III. A prelimi-
eS,
nary account of microfungi colonizing Laurus nobilis
leaf litter. Trans, Br, mycol, Soc, 77: 457-473.
Kirk, P.M. 1983. New or interesting microfungi X. Hyphomyce-
tes on Laurus nobilis leaf litter. Mycotaxon 18: 259-298.
Matsushima, T. 1971. Microfungi of the Solomon Islands and
Papua New-Guinea, Kobe, Japan.
Matsushima, T. 1975. Icones microfungorum a Matsushima lecto-
rum, Kobe, Japan.
Nees von Esenbeck, C.G. 1817. Das System der Pilze und Schwdam-
me, (Ueberblick), 86 pp.
Pa wozynski,,. KoA. 1962. Circinotrichum and (Gynothrix. ‘Mycol.
BaD. eo WDD.
Pirozymemt, K.A. -and"C.s. Hodges; Jr. 1973. New Hyphomycetes
from oOuth Carolina. Cans J. (Bot. 751°:15 7-173.
Saccardo, P.A. 1880. Conspectus generum fungorum Italiae in-
feriorum. Miicheliav2: 135° pp.
Slusconyeb.c. LOSO*Microfungs “from Australian leaf litter.
IDs Reon ig Sete Merely Cabs alee)
GACCONL bu. , Os ONOLlri and A.M. .Perstani. 1984. Hyphomycetes
rari, Oo inberessanti ;della foresta tropicale., Ll.) Pyri-
cularia fusispora comb.nov., nuova combinazione per la
specie Nakataea fusispora, Micol. Ital. 2: 7-10.
N43 oy WE MAR in a es Ae | Re | ,
, wri 4 EA me hail my ot [ ; ¥ he
sith LGC AAA KEANE AAAS a oe
iu oy ‘ee wall’ baseline) i
wk
va a Oe a 5 hl Bey t . ‘ i i a : te, :
} 4 a
ai i
4
i tlh Ht HN “ahi aah) i sh) mau { ey soir
i iy il ene ne, K i f Wien aig? Me it wf whet os nay Hina fi) iy
i wh ; hte Bit Vig diae f eh a “hy ut Wa ag) hae, ays ni aa" Mie heen
4 ve Went ne he 4 i a:
mee Win his f ae ied f
i W rahe! Bre Ly iia su bo idealau tins Pi
i a ra i Payee ares a tae AL vi ey a %
i me Ai tee ah uw, ave ‘ et it rt wall LY been
bith: PO) ee en aie WA ued n ee
PR ia hy | oo mai it) thy, (ae
fig ni i ip ihe ; Wy oe xy at Be hy ae aiiiks aly’ a, ah Any Mea! i
i Bpiae eit ny Hil ik atte sR TE) me \ it
te 4 a Viout ah
i yi Dae a ie
Mii
; La ] 7 ‘
Ly Viet gal ! (erie
“yt q " ; dR Te fae ivy ae vie.
at : ‘han Linn Yo, oi) ie
iV : j PAN Cone A 7] | . mt
i a ' ! Pio
Ws be te aa. BT eiT 7 A on ue 7 | ¥ x
PAP el mre ny Vids Mae: © i ay vs mi y ' Weer allt ren Pe cee ee it y ava
buat A aug nt ; ‘ly rei YY eee R PD hae
OO Sane he er A Te iach (inde As), ‘vO io ,
rR, inc she nie dey ene AA villa i Aa on He dl
Dee a ney " i Wi) vy Lay Whi we aia a hy
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MYCOTAXON
VOleeAAVid , PP eloo~ bg October-December 1986
DIGITATISPORA LIGNICOLA, A NEW MARINE
LIGNICOLOUS BASIDIOMYCOTINA
E.B. GARETH JONES
School of Btologteal Setenees, Portsmouth Polytechntec,
King Henry I Street, Portsmouth, PO1 2DY, England.
While collecting marine fungi on San Juan Island,
Washington State, U.S.A., a new Basidiomycotina with
tetraradiate basidiospores was encountered (Jones, 1985).
This differs markedly from Dtgttattspora martna (Doguet,
1962; Kohlmeyer and Kohlmeyer, 1979), which was also
collected at the same site.
DIGITATISPORA LIGNICOLA sp. nov. (Figs. 1-10, 14-15)
Hyphae 2.5um diametro, hyalinae, fibuligerae. Bastdto-
carpt 30-120um alti 0.5mm lati, 1-2mm longi, irregulares,
resupinati, albi, molles. Aymentwn 30-80um altum, sine
Cy SULdta Ss ADASTOUd rae. ot (45151) 7.0.3 moe (988) On lum,
2-4-spora, clavata, erecta, hyalina, sine sterigmatis.
Bastdtosporae tetraradiatae, hyalinae, ramis primis 13.4
e741 xo. 3) 07.5) 136m, ferectis,) ad: basim atten-
Uatre witanis ‘laceralipus 6.9 \0L4"0) 12900 x 2.ouNo Joyo Oo!
ume ramisevicinis 6.9) (16.6). 2503) xi 2290 (4,0)7. 6. 90m.
radiantibus et attenuatis. Habitat in ligno. Holotypus
IMI 292870.
Hyphae 2.5ym in diameter, septate, with clamp connections,
hyaline. Basidiocarps 30-120um high, 0.5mm wide, 1-2mm
long, irregular, resupinate, white, soft. Hymenium 30-80
(mun enewLonout. cystidias, uBasidiams2. 5) (4ol)vsda Kr Oo
(9.8) 10.lum, 2-4 spored, clavate, erect, hyaline without
sterigmata. Basidiospores tetraradiate, occasionally
with secondary and tertiary branching, hyaline, deciduous,
pasate rinn lo. 0028,.5)* 41 s5um so. ot C7) oum .erect,
broad but tapering to the attachment pore to the basidiun,
apical arms: opposite arm 6.9 (14.6) 23.0 x 2.8 (3.8) 8.1
(im eeadjacent.arms:6.9 °(16.6),25.3) x 2.3; (4.0) "6.9m
radiating and attenuated.
Habitat on driftwood.
Holotype Herb CMI (IMI) 292870
Etymology: Latin ltgnum = wood and cola = dweller in
reference to the substratum of the species.
A Nis
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157
LEGENDS
Pomel On0 toy tate spond witgntcola, 1,2 .-4,).6,
Basidiocarps on wood with club-shaped basidia
(arrowed). 3. Basidium with two basidiospores,
arrow indicates point of attachment. 5. Basidium
with spores and a hypha bearing clamp connections
arrowed. /-10. The tetraradiate hyaline basidio-
spores, and showing branching of one arm. All
interference light micrographs. Bar lines: 20m.
Pigs lial. Yiotrarrepora marina. 11,12. Basidia
bearing tetraradiate basidiospores, point of attach-
ment arrowed. 13. Basidiospore. Figs. 14-15.
Digttattspora ltgntcola. Basidiospore attachment to
basidium arrowed. All interference light micro-
graphs. Bar lines: 20ym.
158
D. ltgnteola resembles D. marina but differs in the
following respects:
D. ltgnteola D. martna
Basidiocarps diffuse irregular but
tufts on wood clearly recognisable
basidiocarps on wood
Basidia club-shaped cylindrical to
short subclavate
BYP S iri [RWG |< 50 x 5-6.5um
6.9-16.1lum
Basidiospores arms not separated radiating arm,
pointed and cylindrical
frequently branched rarely forked
once or twice 26.5-41 (-45) x
18.4-41.5 x 2 Sao
S237 > OumM
The basidia and basidiospores differ in size and
morphology and these are considered to be significant at
the species level. However, further collections and
observations may indicate that these differences are
sufficient to warrant separation at the generic level.
This species brings the number of lignicolous marine
Basidiomycotina to four although other species have been
reported on marine Angiospermae or free floating (yeasts)
(Kohlmeyer and Kohlmeyer, 1979). Of the non-yeast
Basidiomycotina Wia vtbrtissa and Digtitattspora martina
are the most frequently encountered in temperate locations
(Byrne and Jones, 1974; Kohlmeyer and Kohlmeyer, 1979;
Jones, Rule and Jones, 1985). In the tropics Halocyphtna
vtllosa seems to be particularly common on decaying
mangrove wood (Kohlmeyer, 1984; Hyde, 1985; Jones,
unpublished) and is often the dominant species encountered.
ACKNOWLEDGEMENTS
I am grateful to C. Derrick for photographic assistance,
Dr. and Mrs. R.A. Loveless for the Latin diagnosis, the
Director and staff of the Friday Harbor Marine Laboratory,
University of Washington for laboratory facilities and
their help and to the Royal Society for the award of a
travel grant from the Marshall and Orr Bequest.
133
LITERATURE
BYRNE, P.J. and JONES, E.B.G. 1974. Lignicolous marine
fungi. Veroeff. Inst. Meeresforsch. Bremerhaven,
SUD Ye tw oUL=o20.
DOGUET, G. 1962. Dtgitattspora marina, n.g., n.sp.,
Basidiomycete marine. C.R. Hebd. Seances Acad.
SG O44 30-9 50..
HYDE, K. 1985. Spore settlement and attachment in marine
funelr ven, De Kihesas, \CCEN WALA) Portsmouth
Polytechnic.
JONES, E.B.G. 1985. Wood inhabiting marine fungi from
San Juan Island with special reference to ascospore
appendages.) Bot. J. Linn. SOc...) 913 219=231l.
JONES, E.M., RULE, M. and JONES, E.B.G. 1985.
Conservation of timbers of the Tudor Ship Mary Rose.
TnoibiodeterloratLony On) Cbd), io. DALEY) LD. Re
Houghton, G.C. Llewellyn and C.E. O'Rear),
Commonwealth Agricultural Bureaux, London (in press).
KOHLMEYER, J. 1984. Tropical marine fungi. Marine
Ecology 9: (32973/8.
KOHLMEYER, J. and KOHLMEYER, E. 1979. Marine mycology:
The higher fungi. Academic Press, New York, San
Francisco, London, 699p.
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MYCOTAXON
VoD AV LISepp. 7 1ol= 167 October-December 1986
MARINE FUNGI FROM SEYCHELLES. VI.
MASSARINA VELATASPORA
A NEW MARINE ASCOMYCOTINA FROM MANGROVE WOOD
K.D. HYDE! and B.D. BORSE-
pepartment of Btological Setences, Portsmouth Polytechnte
King Henry I Street, Portsmouth, PO1 2DY. England.
* nepartment of Botany, Arts, Commerce and Setence College,
Erandol 425109, India.
During a recent study of the marine fungi of the
Seychelles, several species were collected from the
mangrove ecosystem, which were found to be new to science.
Many of these fungi have recently been described:
Aigtalus grandis, A. parvus(Kohlmeyer and Schatz, 1985),
Antptodera mangrovtt (Hyde, Farrant and Jones, 1986),
Caryosporella rhizophorae (Kohlmeyer, 1985), Halosarpheta
abonnts (Kohlmeyer, 1984), Halosarpheta ratnagirtensts
(Patil and Borse, 1982), Btatrtospora martna (Hyde and
Borse, 1986) and Trematosphaerta ltgnatitlts (Kohlmeyer,
1984). In this paper a new species belonging to the
genus Massarina is described. Massartna velataspora
is characterised by having bitunicate asci, with an apical
apparatus, ascospores which are 1-3 septate, hyaline and
surrounded by mucilaginous sheath. M. velataspora was
collected in Seychelles on 9 occasions and was classified
by Hyde (1986) as occurring, occasionally. It has also
been collected from Maharashtra (India) and Brunei.
MASSARINA VELATASPORA HYDE et BORSE sp. nov. (Figs. 1-8)
Ascocarpt 700-(800)-1200um lati, solitarii vel gregarii,
subglobosi, immersi, erumpesdentes, ostiolati, papillati,
coriaces, brunnei vel compressi, nigri, ascocarpi expositi
hyphis mutis ontecti. Cotza concia, 55-300um longa x
60-200um lata media et nigra. Pseudoparaphyses 1.1-1.7ym
lati, angusti, hyalini, simplices et numerosi. Asct
220-(251)-320 x 23(28.2)-34um, octospori, cylindrici,
parvi, pedunculati, bitunicati, pachydermi, physoclasti,
cum apparatu apicali, ad basim ascocarpi orientes.
Ascosporae 45-(49.5)-56 x 14-(15.9)-19ym, uniseriatae vel
biserliatae, fusiformatae vel ellipsoidales, 1-3 septatae,
hyaline, constrictione ad septum, cum cellis mediis
maioribus, cellis, quando adsunt polaribus parvis et
elongatis, ascosporae mucilage circumvelatae.
162
i)
Figs. 1-3 Massarina velataspora: light micrographs. id De
Ascospores, 1-3 septate, hyaline, and surrounded by an
indistinct mucilaginous sheath (2-arrowed). 3. Squash
showing ascospores and pseudoparaphyses (arrowed - 8).
Ascospore end cells are small and elongate. Bar lines =
10um.
163
Figs. 4-6 Massartna velatospora: light micrographs.
Asci. Note the apical chamber (arrowed in 5). Spores
released by the rupture of the ectoascus (EC) and the
stretching of the endoascus (En). Bar lines = 10pm.
164
Figs. 7, 8. Massartna velatospora: Scanning electron
microscope micrographs. 7. Ascospores settled on a poly-
carbonate membrane - mucilaginous sheath extends over
pores (arrowed) in the membrane. 8. Ascus tip surrounded
by filamentons pseudoparaphyses. Bar lines = 10pm.
165
Colontae in agaro cum corn-mealo et aqua maris composito
floccosae crescentes, albae, purpureae maturescentes, et
agarum colorantes purpuram. Structuras similes pycidio-
rum formantes quae maturescentes non visae sunt.
Substratum lignum mortuum. Dtstrtbutto Oceanus Indicus
(Seychelles, India), Mare Chinese Austrum (Brunei).
Ascocarps 700-(870)-1200um in diameter, immersed,
becoming rumpent, subglobose or depressed, ostiolate,
papillate, coriaceous, brown to black, solitary or gre-
garious, exposed tissue covered in a fine hyphal mat.
Necks 55-300yum high x 60-200um in diameter, central and
black. Pseudoparaphyses 1.1-1.7um in diameter, filament-
ous, hyaline, simple and numerous. Asc? 220-(251)-320 x
23-(28.2)-34um, 8-spored cylindrical, short pedunculate,
bitunicate, thick-walled, with an apical apparatus, and
developing at the base of the ascocarp venter.
Ascospores 45-(49.5)-56 x 14-(15.9)-19ym, uni-seriate,
fusiform, to ellipsoidal, 1-3 septate, hyaline, constrict-
ed at the septa, central cells larger, apical cells when
present small and elongate, surrounded by an indistinct
mucilaginous sheath, 5-22ym wide. Colonies on sea water
cornmeal agar, floccose, at first white becoming purple
with age, and staining the medium purple. Forming
pycnidial-like structures, which were not observed to
mature. Substrate intertidal mangrove roots and branches
of Rhizophora mucronata. Range. Indian Ocean (Seychelles,
India) and South China Sea (Brunei). Type material:
January 1984, Herb. IMI 297770, slides 1-10 (Holotype)
and dried wood (Isotype).
Etymology: From the Latin, velata = a wrapping or enve-
lope, in relation to the envelope of mucilage sheathing
the spore and spora = spore. The species belongs in the
Pleosporales, Massarinaceae, and in the genus Massarina
as it has fusiform-ellipsoidal ascospores, which are
hyaline, 1-3 septate and surrounded by a mucilaginous
sheath (Figs. 1-3). Aset are 8-spored, cylindrical and
bitunicate (Figs. 4-6). A number of aquatic Massarina
species have been reported: M. cystophorae (Cribb et
Herbert) Kohlm. et Kohlm. (Kohlmeyer and Kohlmeyer, 1979);
M. aquattca Webster (Webster, 1965); Massarina sp.
(Webster and Descals, 1979). M. velatospora differs from
the only known marine species, M. cystophorae in spore
size, the latter only has cap-like gelatinous appendages
and is parasitic on the brown algae Cystophora retroflexa
and C. subfarctnata.
166
Measurements of aquatic Massarina sp.(um).
Ascocarps' Asci Ascospores
M.cystophorae 650-950 x 105-150 x 50-65 x
650-1000 50-64 16-23
M. velatospora),),/00=1200))) | 220-3205 (i 45—56) x
23-34 14-19
M. aquatica 300-500 TLO=130 °x eos o0ex
10-1205 Dae
Massartna sp.1 120-240 96-120 hare eae
9-10 1-9
Massartna sp.2 140-260 68-82 x 20-24 9x
TZ oa)
Many marine fungi have been shown to have mucilaginous
sheaths: Leptosphaerta contecta Patouillard, Pleospora
guadefroyt Kohlm. (Kohlmeyer and Kohlmeyer, 1979), and
Nimbospora btpolarts Hyde et Jones (Hyde and Jones, 1985).
Of the marine fungi associated with mangroves, a mucila-
ginous sheath has also been noted in Leptosphaerta
avicenntae Kohlm. et Kohlm. (Kohlmeyer and Kohlmeyer,
1979). Hyde, Jones and Moss, (1986a,b), have suggested
that mucilaginous sheaths aid in the attachment of spores
to surfaces. Fig./7 shows the mucilaginous sheath of
M. velatospora which is amorphous, almost electron trans-
parent and closely adpressed to the polycarbonate
membrane. Undoubtedly the presence of a sheath increases
the surface area of the spore and thus enhances the spore
attachment. Similar observations have been made for
Leptosphaerta contecta, spores becoming readily attached
to glass slides. The sheath also causes the ascospore to
float on the surface of water, the sheath "sticking" to
the surface film on the water. This may also be advanta-
geous to the ascospore, which will float on the incoming
and outgoing tides, or be wind blown and be dispersed in
this manner.
ACKNOWLEDGEMENTS
K.D. Hyde is grateful for the award of an SERC student-
ship, and also to CIBA-GEIGY A/S, Basle, for financial
support to visit the Seychelles. We are also indebted
to C. Derrick for photographic assistance, P. Crook for
assistance in the Seychelles, Mrs. Monica Heywood-Kenny
for the Latin diagnosis and to Professor E.B.G. Jones
for his encouragement and also reviewing the manuscript.
167,
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HYDE, K.D. 1986. Frequency of occurrence of lignicolous
marine fungi in the tropics. In: "Biology of Marine
Fungi" (Ed. S.T. Moss), pp.311-322, Cambridge,
Cambridge University Press.
HYDE, K.D. and BORSE, B.D. 1986. Marine fungi from
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HYDE, K.D. and JONES, E.B.G. 1985. Marine fungi from
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HYDE, K.D., FARRANT, C. and JONES, E.B.G. Marine fungi
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MYCOTAXON
Vol. XXVII>" pp.” 169-182 October-December 1986
TAXONOMIC CONCEPTS IN THE ENDOGONACEAE: III. THE
SEPARATION OF SCUTELLOSPORA GEN. NOV. FROM GIGASPORA
GERD. & TRAPPE
CHRISTOPHER WALKER
Forestry Commission, Northern Research Station, Roslin, Midlothian, EH25 9SY,
UK
and
F. E. SANDERS
Department of Plant Sciences, University of Leeds, Leeds, LS2 9JT,
UK
SUMMARY
The genus Gigaspora is split into two genera, Gigaspora Gerd. & Trappe emend. Walker &
Sanders and Scutellospora gen. nov. This separation is based principally on details of spore
germination, although other characteristics are also used. The factors that characterise and
separate the two genera are detailed.
The genus Gigaspora Gerd. & Trappe was erected to contain fungi in the
Endogonaceae which form arbuscular mycorrhizas and produce spores on a bulbous
suspensor-like cell (Gerdemann & Trappe 1974) (Fig. 1). The genus originally
included five species, and a further 18 have since been added (Becker & Hall 1976;
Hall 1977; Nicolson & Schenck 1979; Ferrer & Herrera 1981; Bhattacharjee et al.
1982; Schenck & Smith 1982; Koske, Miller & Walker 1983; Hall & Abbott 1984;
Koske & Walker 1984, 1985). Observations of spore wall structure, germination
characteristics of the spores, and ornamentation of auxiliary cells (terminology of
Trappe & Schenck (1982)), show that there are two distinctive groupings within the
genus (Walker 1985; Walker & Sanders 1985). These differences form a clear and
consistent dichotomy of Gigaspora sensu lato and it is appropriate to assign each of
the groupings to a different genus.
The term azygospore, used by some authors for the large spores of Gigaspora carries
with it the connotation of asexuality. While we do not consider there is, as yet,
evidence for a sexual process in the formation of these spores, neither do we know of
any evidence to indicate that they are indeed asexual. Thus we prefer to avoid the use
of azygospore, choosing simply to refer to them as spores. This is in agreement with
the usage of Koske et al. (1983), where the use of the term was questioned in the
description of G. reticulata Koske, Miller & Walker, and of Koske & Walker (1984,
1985), where it was dropped without comment.
170
Fig. 1: Spores of species of Gigaspora Gerd. & Trappe. Note the typical bulbous suspensor-
like cell upon which the spore is borne.
a) Intact spore of G. margarita, a species with only a single wall group, lacking the flexible
inner walls illustrated in Fig. lc & d.
b) Accrushed spore of G. margarita. Note the lack of membranous inner walls.
c) Aspecies of Gigaspora sensu lato (G. heterogama) that possesses an inner wall group with
flexible inner walls, detectable in this intact spore by the plasmolysis at upper right.
d) This crushed specimen of an undescribed species of Gigaspora sensu lato has flexible inner
walls (small arrows) forming an inner wall group. The spore also has a distinct germination
shield (gs).
171
On another matter of terminology, in the literature on the Endogonaceae, the term
‘spore’ has generally been used in a rather broad and uncritical manner. For example,
the so-called ‘spores’ of members of the genus Endogone are, in reality,
zygosporangia enclosing a zygospore (Berch & Fortin 1982). Whether the ‘spores’ of
species of Gigaspora sensu lato should be considered also to be sporangia (as
suggested by Gibson (1985)) enclosing a single spore remains moot, but those herein
placed in the new genus Scutellospora do seem to produce a definite endospore.
Nevertheless, we shall retain the traditional usage of the term ‘spore’ in this paper.
Wall structure
The terminology of Walker (1983, 1986) will be used in discussion of spore wall
characteristics. Spores formed by species of Gigaspora sensu lato can be classified
into one of two types, those that have one or more flexible inner walls and those that
do not. This dichotomy can be inferred from the original descriptions in Nicolson &
Gerdemann (1968), though these authors made no comment on the matter. Spores of
species such as G. gigantea (Nicol. & Gerd.) Gerd. & Trappe have only a single wall
group which usually consists of an outer, unit wall, tightly adherent to an inner,
laminated wall. Although they must have a plasmalemma, they do not possess
flexible inner walls, either membranous or coriaceous. In contrast, although spores
of species typified by G. calospora (Nicol. & Gerd.) Gerd. & Trappe may have an
outer wall structure similar to that of G. gigantea, they invariably also have at least
one inner wall group, consisting of at least one membranous or coriaceous wall. This
inner wall group seems to form a complete endospore that becomes detached at the
spore base at maturity, and which usually can be readily separated from the other
walls. Both spores of the G. gigantea type and those of the G. calospora group share
the possession of a thin inner wall layer, adjacent to the plasmalemma, which is
approximately 0.07 um in thickness, and thus is not resolvable by light microscopy.
This layer is attached to the outer wall group in the former, and the inner wall of the
innermost wall group in the latter (Gibson 1985).
Germination characteristics
The germination characteristics of spores in Gigaspora sensu lato are linked closely
with the differences in wall structure. Spores of the G. gigantea group germinate by
production of one or more germ tubes directly through the spore wall, normally at a
point near the suspensor-like cell’s attachment, after the formation of a warty ring on
the interior of the laminated wall (Fig. 2). This mode of germination, discussed and
illustrated for G. margarita Becker & Hall by Sward (1981), will be referred to as
direct germination. The G. calospora group has an entirely different, and much more
complicated method of germination. The germ tube emerges through the outer wall
in amanner superficially similar to direct germination, but it does not originate from
the outer wall group. It emerges from a specialised structure formed within one of the
flexible walls in the inner group, the development of which is not fully elucidated.
Light-microscope studies and examination of whole and crushed or dissected spores
of several species and examination of electron micrographs of Gigaspora heterogama
(Nicol. & Gerd.) Gerd. & Trappe (Gibson 1985) leads us to infer the following mode
of formation (Fig. 3). Initially, a hole appears in a wall of the inner wall group,
through which the cell plasma membrane protrudes. The extruding membrane then
172
Fig. 2: Direct germination in spores of Gigaspora sensu lato.
a) Two germ tubes emerging directly through the spore wall of a G. margarita spore.
b) Scanning electron micrograph (SEM) of the inside of a spore with direct germination. The
point of attachment of the suspensor-like cell can be seen at upper right (large arrow). Four
sites of germ tube initiation can be seen at lower left (small arrows).
c) Detail of two of the germ tube initiation points in Fig. 2b. Note the build-up of wall
material to form a collar (C) around the germ tube initial (gti).
d) Germ tube initial in cross section by light microscopy, showing build-up of collar material
around the lumen of the germ tube initial. The relatively simple wall structure of a unit wall (1)
and a laminated wall (2) in a single group (A) is illustrated.
17S
Y
A
Fig. 3: Diagrammatic representation of the stages in germination shield formation (in plan
view). The figure should be viewed as if the paper were a flexible (membranous or coriaceous)
inner wall and the outer wall group was not present. The arrows indicate direction of growth
and folding of the cell wall as the next stage of development proceeds.
a) A hole appears in the flexible wall.
b) The plasma membrane extrudes and extends, sandwiched between inner and outer wall
groups.
c&d) Growth and development takes place as the two lobes of the germination shield are
formed.
e) Inthesimplest form of germination shield, the two lobes meet and a germ tube initial (gti)
is formed in each lobe. Wall material is deposited to complete the germination shield.
f) In amore complex germination shield, further lobes are formed which meet and from
which germ tubes may also emerge. Yet more complex forms with several lobes can be found in
some species.
174
\
Fig. 4: Light micrographs of germination shields in two species of Gigaspora sensu lato.
a & b) Plan views of the simplest form of germination shield from spores of Gigaspora
calospora. The lobe on the right side is damaged and somewhat displaced in a. A germ tube is
emerging from the lobe on the right in b.
c&d) Planviews of complex forms of germination shield with several lobes; c. is G. nigra and
d. is an undescribed species from Nigeria.
175
folds back on itself to produce a characteristic ‘wishbone’ formation. The lobes thus
formed meet in a pincer-like configuration reminiscent in plan view of the suspensor
cells of a zygospore, but without the spore between them. The structure thus formed
is constrained by its juxtaposition between the inner and outer wall groups, into the
shape of a convex, more or less discoid shield. Additional wall material is then
deposited to encompass this shield, thickening around the edges and folds. In section,
the shield then appears to be formed within the membranous wall. This material
appears to be the same as that of the membranous wall in some species, but in others
may be brittle. In such instances, it may also be pigmented. Two germ tube initials
then form, one on each side of this pincer-like arrangement. A germ tube then arises,
often from only one of these initials, and emerges through the outer spore walls.
In its simplest form (found, for example on G. calospora and G. heterogama), there is
no further division (Fig. 4a), but in some species, new lobes form repeatedly, dividing
the structure into several compartments, each with a germ tube initial. An example of
this complex type of structure is illustrated by G. nigra Redhead (Fig. 4b). The phrase
‘germination compartment’ (e.g., Hall 1984) has been used for this structure, but
Koske et al. (1983) and Koske & Walker (1985) termed it a ‘germination shield’.
Figures 5 & 6 illustrate the ‘compartment-like’ features which prompted the use of the
former term. We prefer the use of the latter phrase because it is descriptive of both
form and function, rather than merely of the form viewed in optical section (Hall
1977). Only one germination shield is formed in each spore, though it may produce
more than one germ tube.
Fig. 5: Light micrograph of an optical section through the germination shield of Gigaspora
calospora. The hole created at initiation of the shield can be seen (h), and the two lobes which
form the compartments (c) from which germ tubes may emerge are delimited by the folded
membranous wall (m). The fold in the membranous wall is out of focus on the right of the hole.
heel
PLAN
—ae-
outer
wall
SECTION (X--X)
Fig. 6: Diagrammatic representation of the simplest form of germination shield (as in
Figs. 4a & 5) drawn as a plan view and a cross section through the line X-X which passes
through the hole (h) that appears at the initiation of shield formation. This form of shield has
only two lobes each of which creates a compartment (c) with a germ tube initial (gti). A germ
tube is shown emerging from the lobe on the right (gt).
177
The original descriptions of G. calospora and G. gigantea did not distinguish between
their germination modes, indicating merely that both species germinated directly
through the spore wall near the attachment. This was discussed by Hall and Abbott
(1984) who, from examination of the type material of G. calospora, showed that its
germination was by means of a germination shield.
Germination shields may have taxonomic value at species level in the future and they
should be described and illustrated whenever possible in species descriptions.
Auxiliary cells
When the first species of the group were formally described (Nicolson & Gerdemann
1968) it was recognised that auxiliary cells were of two forms (Fig. 7). Both begin as
smooth sacs (Fig. 7a), but as they mature, they develop different kinds of
ornamentation. The first type, represented by G. gigantea, is decorated by
echinulations or narrow papillae, which may be cut off from the main body of the sac
by septa (Fig. 7b). The other form, of which G. calospora is typical, has auxiliary cells
that become knobby due to the formation of much broader, blunt projections or very
broad, relatively low papillae (Fig. 7c). Although there may be some minor
differences among species (Trappe 1982), such differences are merely variations of
these two basic forms. Examination of the literature and of specimens of most of the
currently described species indicates that each has exclusively one or other of these
types. Though Nicolson & Gerdemann (1968) indicated that the auxiliary cells of
G. heterogama were smooth, they are in fact knobby (Koske & Walker 1985), and it is
probable that the former authors were observing immature cells. Regardless of minor
variations, therefore, these will be termed ’echinulate’ and ’knobby’ auxiliary cells
(Figs. 7 & 8). The species with spores that have only a single wall group and that
germinate by direct germination have the former, while the species with membranous
inner walls that germinate by means of a germination shield have the latter. The shape
a b Cc
Fig. 7: Drawings of auxiliary cells in Gigaspora sensu lato.
a) Immature auxiliary cells before acquisition of ornamentation.
b) Echinulate auxiliary cells typical of the G. gigantea group within the genus.
c) Knobby auxiliary cells, typical of the G. calospora group.
178
of auxiliary cells has been thought to have some taxonomic value at the specific level
(Trappe 1982), but Koske & Walker (1985) give this character little weight because of
the large variation within, and the similarities among species.
Summary of characteristics separating the two groupings
The genus Gigaspora Gerd. & Trappe can be divided into two distinctive genera,
primarily on the basis of germination characteristics, but also of spore wall structure
and morphology of auxiliary cells. In one group, for which the generic name
Gigaspora will be retained in accordance with the International Code of Botanical
Nomenclature (Article 52), germination is by egress of a germ tube directly through
the outer spore wall, spores do not possess inner wall groups of (flexible)
Fig. 8: The two types of auxiliary cells in Gigaspora sensu lato.
a&b) Echinulate auxiliary cells from an undescribed species, and G. margarita, respectively.
c&d) Knobby auxiliary cells from an undescribed species and G. heterogama, respectively.
179
membranous or coriaceous walls, and auxiliary cells are echinulate. This group is
exemplified by the species Gigaspora gigantea. The second group, represented by
Gigaspora calospora, has spores which germinate by means of a germination shield
formed in an inner wall group of membranous or coriaceous walls. Auxiliary cells of
these fungi are knobby. Many species in this group have ornamented outer walls
(Koske & Walker 1985), whereas no described species in the first group does.
Auxiliary cells are delicate and not always easy to find, except from pot cultures.
Germination shields form at a late stage in the development of the spore, may not be
present in all specimens, and may be very difficult to see on species with hyaline
spores. Nevertheless, the presence or absence of an inner wall group with at least one
flexible wall can always readily be detected, and a species can be placed in its group on
this characteristic alone. There thus will be no difficulty in identifying the appropriate
grouping for a species. The G. calospora grouping is herein given separate generic
status as Scutellospora gen. nov.
GIGASPORA Gerdemann & Trappe (Mycologia Memoir No. 5, p. 25, 1974) emend.
Walker & Sanders
Spores produced singly in soil, large variable in shape, usually globose or subglobose,
but often ovoid, obovoid, pyriform, or irregular, especially when constrained during
formation, borne on a bulbous suspensor-like cell, usually with a narrow hypha
extending from one or more peg-like projections towards the spore. Spore wall
structure of a single wall group, lacking flexible (membranous or coriaceous) walls.
One or more germ tubes produced directly through the spore wall near the base.
Thin-walled, echinulate or finely papillate auxiliary cells borne in soil, on straight or
coiled hyphae, formed singly or in clusters. Forming endomycorrhizas with
arbuscules and hyphal coils, but without vesicles.
SPECIES INCLUDED
Gigaspora gigantea (Nicol. & Gerd.) Gerd. & Trappe, Mycologia Memoir No. 5,
pp. 29-30. 1974.
= Endogone gigantea Nicol. & Gerd., Mycologia 60: 321. 1968.
Gigaspora margarita Becker & Hall, Mycotaxon 4:1-2. 1976.
Gigaspora candida Bhattacharjee et al., Transactions of the British Mycological
Society 78:184. 1982.
Gigaspora decipiens Hall & Abbott, Transactions of the British Mycological Society
83:204. 1984.
Gigaspora rosea Nicol. & Schenck, Mycologia 71:190. 1979.
Gigaspora albida Schenck & Smith, Mycologia 74:85. 1982.
SCUTELLOSPORA Walker & Sanders, gen. nov.
Sporae in solo singulariter productae, terminaliter in cellula bulbosa suspensoriformi portatae, plerumque
hypha angusta e cellula suspensoriformi ad sporam extensa. Paries sporae stratosus, stratis in turmis
duabus vel ultra, turma interiore stratorum flexibilium composita. Scutum germinationis prope basim
sporae in vel inter stratis flexibilibus interioribus formatum, tubo germinationi uno vel ultra ex scuto
180
germinationi formato. Cellulae auxiliares nodosae singulariter vel fasciculatae in solo portatae. Fungus
endomycorrhizas cum arbusculis et hyphis spiralibus sed sine vesiculis formans. Typus - Scutellospora
calospora (Nicolson & Gerdemann) Walker & Sanders.
Spores produced singly in soil (or rarely in cortical cells of roots), large, variable in
shape, usually globose or subglobose, but often ovoid, obovoid, pyriform, or
irregular especially when constrained during formation; borne on a bulbous
suspensor-like cell, usually with a narrow hypha extending from one or more peg-like
projections towards the spore. Spore wall structure of at least two wall groups, with
one or more flexible membranous or coriaceous walls in the inner group or groups.
Germination by means of one or more germ tubes produced near the spore base from
a germination shield formed upon or within a flexible inner wall. Thin-walled,
knobby or broadly papillate auxiliary cells borne in soil, on straight or coiled hyphae,
formed singly or in clusters.
Forming endomycorrhizas with arbuscules and hyphal coils, but without vesicles.
ETYMOLOGY: Latin, scutellum, small shield and spora, spore; referring to the
production of germination shields in spores of members of the genus.
SPECIES INCLUDED
Scutellospora calospora (Nicol. & Gerd.) Walker & Sanders, comb. nov.
=Endogone calospora Nicol. & Gerd., Mycologia 60:322. 1968.
=Gigaspora calospora (Nicol. & Gerd.) Gerd. & Trappe, Mycologia Memoir
No. 5. pp. 28-29. 1974.
Scutellospora heterogama (Nicol. & Gerd.) Walker & Sanders, comb. nov.
=Endogone heterogama Nicol. & Gerd., Mycologia 60: 319-320. 1968.
= Gigaspora heterogama (Nicol. & Gerd.) Gerd. & Trappe, Mycologia Memoir
No. 5. p. 31. 1974.
Scutellospora alborosea (Ferr. & Herr.) Walker & Sanders, comb. nov.
= Gigaspora alborosea Ferr. & Herr., Rev. Jardin Bot. Nac., Habana 1:55-57.
1981.
Scutellospora minuta (Ferr. & Herr.) Walker & Sanders, comb. nov.
= Gigaspora minuta Ferr. & Herr., Rev. Jardin Bot. Nac., Habana 1:53-54. 1981.
Scutellospora aurigloba (Hall) Walker & Sanders, comb. nov.
= Gigaspora aurigloba Hall, Transactions of the British Mycological Society
68:351. 1977.
Scutellospora savannicola (Herr. & Ferr.) Walker & Sanders, comb. nov.
= Gigaspora savannicola Herr. & Ferr. in Ferr. & Herr., Rev. Jardin Bot. Nac.,
Habana 1:57-60. 1981.
Scutellospora tricalypta (Herr. & Ferr.) Walker & Sanders, comb. nov.
= Gigaspora tricalypta Herr. & Ferr. in Ferr. & Herr., Rev. Jardin Bot. Nac.,
Habana 1:49-51. 1981.
181
Scutellospora erythropa (Koske & Walker) Walker & Sanders, comb. nov.
= Gigaspora erythropa Koske & Walker, Mycologia 76: 250-252. 1984.
Scutellospora dipapillosa (Walker & Koske) Walker & Sanders, comb. nov.
= Gigaspora dipapillosa Walker & Koske in Koske & Walker, Mycologia
77:709-710. 1985.
Scutellospora persica (Koske & Walker) Walker & Sanders, comb. nov.
= Gigaspora persica Koske & Walker, Mycologia 77:708-709. 1985.
Scutellospora verrucosa (Koske & Walker) Walker & Sanders, comb. nov.
= Gigaspora verrucosa Koske & Walker, Mycologia 77:705-706. 1985.
Scutellospora reticulata (Koske, Miller & Walker) Walker & Sanders, comb. nov.
= Gigaspora reticulata Koske, Miller & Walker, Mycotaxon 16:429-433. 1983.
Scutellospora gregaria (Schenck & Nicol.) Walker & Sanders, comb. nov.
= Gigaspora gregaria Schenck & Nicol in Nicol. & Schenck, Mycologia 71:185-
186. 1979.
Scutellospora pellucida (Nicol. & Schenck) Walker & Sanders, comb. nov.
= Gigaspora pellucida Nicol. & Schenck, Mycologia 71:189-190. 1979.
Scutellospora nigra (Redhead) Walker & Sanders, comb. nov.
= Gigaspora nigra Redhead in Nicol & Schenck, Mycologia 71:187-188. 1979.
Scutellospora gilmorei (Trappe & Gerd.) Walker & Sanders, comb. nov.
= Gigaspora gilmorei Trappe & Gerd. in Gerd. & Trappe, Mycologia Memoir
No. 5. pp. 27-28. 1974.
Scutellospora coralloidea (Trappe, Gerd. & Ho) Walker & Sanders, comb. nov.
= Gigaspora coralloidea Trappe, Gerd. & Ho in Gerd. & Trappe, Mycologia
Memoir No. 5. pp. 30-31. 1974.
ACKNOWLEDGEMENTS
We are grateful to Dr. R. E. Koske, University of Rhode Island, and Dr. N. C. Schenck, University of
Florida for kindly reviewing the manuscript and for their helpful comments, and Dr. Bud Uecker,
Mycology Laboratory, Beltsville, Maryland for preparing the Latin diagnosis. We also thank Colin
McEvoy, Forestry Commission for help in preparing the plates.
REFERENCES
Becker, W. N. and Hall, I. R. 1976. Gigaspora margarita , anew species in the Endogonaceae. Mycotaxon
4:155-160.
Berch, S. M. and Fortin, J. A. 1982. Germination of zygospores of Endogone incrassata. Mycologia
74:86 1-864.
Bhattacharjee, K. G., Mukerji, K. G., Tewari, J. P. and Skoropad, W. P. 1982. Structure and
hyperparasitism of a new species of Gigaspora. Transactions of the British Mycological Society
78: 184-188.
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Ferrer, R. L., and Herrera, R. A. 1981. El genero Gigaspora Gerdemann et Trappe (Endogonaceae) en
Cuba. Rev. Jardin Bot. Nacional, Habana 1(1):43-66.
Gerdemann, J. W. and Trappe, J. M. 1974. The Endogonaceae in the Pacific Northwest. Mycologia
Memoir No. 5. 76 pp.
Gibson, J. L. 1985. Morphology, cytology and ultrastructure of selected species of Endogonaceae
(Endogonales: Zygomycetes). Unpublished PhD dissertation, University of Florida, Gainesville,
Florida. 166 pp.
Hall, I. R. 1977. Species and mycorrhizal infections of New Zealand Endogonaceae. Transactions of the
British Mycological Society. 68:341-356.
Hall, I. R. 1984. Taxonomy of VA mycorrhizal fungi. pp. 57-94. In: VA Mycorrhiza. Ed. by C. LL. Powel
and D. J. Bagyaraj. CRC Press, Florida.
Hall, I. R. and Abbott, L. K. 1984. Some Endogonaceae from South Western Australia. Transactions of
the British Mycological Society, 83:203-208.
Koske, R. E., Miller, D. D. and Walker, C. 1983. Gigaspora reticulata: anewly described endomycorrhizal
fungus from New England. Mycotaxon 16:429-435.
Koske, R. E. and Walker, C. 1984. Gigaspora erythropa, a new species forming arbuscular mycorrhizae.
Mycologia 76:250-25S.
Koske, R. E. and Walker, C. 1985. Species of Gigaspora (Endogonaceae) with roughened outer walls.
Mycologia 77:702-720.
Nicolson, T. H. and Gerdemann, J. W. 1968. Mycorrhizal Endogone species. Mycologia 60:313-325.
Nicolson, T. H. and Schenck, N. C. 1979. Endogonaceous mycorrhizal endophytes in Florida. Mycologia
71:178-198.
Schenck, N. C. and Smith, G. S. 1982. Additional new and unreported species of mycorrhizal fungi
(Endogonaceae) from Florida. Mycologia 74:77-92.
Sward, R. J. 1981. The structure of the spores of Gigaspora margarita. II. Germ-tube emergence and
growth. New Phytologist 88:667-673.
Trappe, J. M. 1982. Synoptic keys to the genera and species of zygomycetous mycorrhizal fungi.
Phytopathology 72:1102-1108.
Trappe, J. M. and Schenck, N. C. 1982. Taxonomy of the fungi forming endomycorrhizae. A. Vesicular-
arbuscular mycorrhizal fungi (Endogonales). Pp. 1-9. In: Methods and Principles of Mycorrhizal
Research. Ed. by N. C. Schenck. The American Phytopathological Society, St. Paul, Minnesota.
Walker, C. 1983. Taxonomic concepts in the Endogonaceae: spore wall characteristics in species
descriptions. Mycotaxon 18:443-455.
Walker, C. 1985. Taxonomy of the Endogonaceae. Pp. 193-199. In: Proceedings of the 6th North
American Conference on Mycorrhizae. Ed. by R. Molina. Forest Research Laboratory, Corvallis,
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Walker, C. 1986. Taxonomic concepts in the Endogonaceae. II. A fifth morphological wall type in
endogonaceous spores. Mycotaxon 25:95-99.
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- Corvallis, Oregon.
MYCOTAXON
Vol. XXVII, pp. 183-192 October-December 1986
ple BE TER 2 SA ea kha ION asthe a ell a Re TIN a i A
FIVE NEW SPECIES OF PARMELIA (LICHENES, PARMELIACEAE)
FROM SOUTHERN AFRICA.
FRANKLIN A. BRUSSE.
Botanical Research Institute,
Private Bag X101, PRETORIA,
SOUTH AFRICA.
ABSTRACT
Five new species of Parmelia (Lichenes, Parmeliaceae) are described from southern
Africa. They are: Parmelia eximia, P. fausta, P. insignis, P. inuncta, and P. pudens.
Their chemistries and affinities are discussed.
PARMELIA EXIMIA Brusse, sp. nov.
Thallus foliosus, saxicola, usque ad 4 cm diametro. Lobi elongati, circa 1.5 mm lati,
120—200 wm crassi. Thallus superne viridis, nitidus, emaculatus, isidiis sorediisque
nullis. Cortex superior 15 ym crassus. Stratum gonidiale 30—50 um crassum, algis
Trebouxiis, 4.5—18 ym diametris. Medulla albida, 80—120 um crassa. Cortex inferior
circa 10 um crassus. Thallus inferne pallidus. Rhizinae 30—60 um crassae. Apothecia
usque ad 5 mm diametris. Hypothecium 35—40 wm crassum. Subhymenium 10—15
um crassum. Hymenium 40—60 ym altum, J+ caeruleum. Asci clavati, cum tholis J+
caeruleis (fig. 1), poricidales. Ascosporae octonae, subsphaericae, 7.0—7.5 x 8.0—8.5
um. Pycnidia globosa, circa 150 um diametris. Pycnidiosporae bacillares, hyalinae,
6—8.5 x 1 um. Thallus acidum usnicum et materias ignotas continens.
TYPUS: SOUTH AFRICA, Cape Province, 65 km WNW of Willowmore, Kouka
river basin, 1 km SE of Rietkuil farmstead, on shale outcrops on gentle S slope, alt.
810 m, F. Brusse 4840, 5 .ii.1986, (PRE, holo-; BM, COLO, LD, US, iso-), fig. 6.
Thallus foliose, saxicolous, up to 4 cm diam. Lobes elongate, about 1.5 mm broad,
120—200 wm thick. Upper surface green, nitid, emaculate, isidia and soredia absent.
Upper cortex 15 ym thick. Algal layer 30—S0 um thick, algae Trebouxia, 4.5—18 um
diam. Medulla whitish, 80—120 um thick. Lower cortex around 10 um thick. Lower
surface pale, rhizinate. Rhizines 30—60 um thick. Apothecia up to 5 mm diam. Hypo-
thecium 35—45 yum thick. Subhymenium 10—15 pm thick. Hymenium 40—60 ym
high, J+ blue. Asci clavate, tholus J+ blue (fig. 1), poricidal with some extension of the
ascus, 8 spored. Ascospores subsphaerical, 7.0—7.5 x 8.0—8.5 um. Pycnidia globose,
about 150 um diam. Pycnidiospores hyaline rods (with 2 subterminal swellings each),
6—8.5 x 1 yum. Chemistry: Usnic acid in the upper cortex, and six unknowns in the
medulla (FB—1 to FB—6, table 1).
184
FIG. 1. Parmelia eximia Brusse, ascus and paraphyses. F. Brusse 4840, holotype. Bar = 10 pam.
Parmelia eximia is here assigned to Xanthoparmelia, but is unique in this section in
possessing subsphaerical ascospores and an unusual chemistry. The subsphaerical asco-
spores are very similar to those of Parmelia sphaerospora Nyl., but the latter is a larger
corticolous species of humid areas, containing atranorin in the upper cortex and a
yellow pigment in the medulla, with or without stictic or hypostictic acids and assoc-
iated compounds, or sometimes gyrophoric acid (Hale, 1976). The unknown substan-
ces in P. eximia appear to be depsidones, as several spots remain unchanged on subjec-
tion to acid hydrolysis. The Rf values of these substances are listed in table 1. Thin
layer chromatography was carried out on the acetone extract of cleaned lobes of these
lichens, in accordance with the method of Culberson (1972), with the modified sol-
vent B, using fert.-butyl methyl ether instead of diethyl ether (Culberson & Johnson,
1982).
Thus far, this unique Xanthoparmelia has only been found in the Kommandokraal
area of the Kouka river basin, some 60 km WNW of Willowmore, in the southern Cape
Province.
185
SOUTH AFRICA, Cape Province — 3322 (Oudtshoorn): 62 km WNW of Willow-
more (2 km WNW of Vlieékraal farmstead), on sandstone outcrops on gentle S slope,
altitude 810 m (—BB), F. Brusse 4844, 5.ii.1986 (PRE, COLO, LD).
PARMELIA FAUSTA Brusse, sp. nov.
Thallus minute foliosus, saxicola, usque ad 4 cm diametro.Lobi sublineares, 0.2—1.0
mm lati, 110—150 um crassi, Thallus superne cinereus, nitidus, emaculatus, isidiis
sorediisque nullis. Cortex superior 15—20 wm crassus. Stratum gonidiale 30—45 um
crassum, algis Trebouxiis 5—15 um diametris. Medulla aurantiaca, 45—90 um crassa.
Cortex inferior 15—20 um crassus. Thallus inferne piceus. Rhizinae simplices, 90—100
lum crassae. Apothecia usque ad 1.2 mm diametris. Hypothecium 30—45 ym crassum.
Subhymenium 35 wm crassum. Hymenium 70 ym altum, J+ caeruleum. Asci clavati,
cum tholis J+ caeruleis (fig. 2), poricidales. Ascosporae octonae, ellipsoideae, 10—12 x
6.5—7.5 pm. Pycnidia ignota. Thallus atranorinum, acidum lecanoricum, et duo
pigmenta ignota (ut in Parmelia schenckiana Mull. Arg.) continens.
TYPUS: SOUTH AFRICA, Cape Province, 19 km § of Prince Albert, summit of
Swartberg Pass, wind blown SE gulleys, on S faces of TMS rock near the ground, alti-
tude 1650 m, F. Brusse 4891, 7.ii.1986 (PRE, holo-; BM, COLO, LD, MEL, US, iso-),
fig. 3.
Thallus minutely foliose, saxicole, up to 4 cm across. Lobes sublinear, 0.2—1.0 mm
broad, 110—150 um thick. Upper surface grey, nitid, emaculate, not isidiate and not
sorediate. Upper cortex 15—20 um thick. Algal layer 30—45 ym thick, algae Trebouxia,
5—15 wm diam. Medulla orange, 45—90 um thick. Lower cortex 15—20 um thick.
Lower surface black. Rhizines simple, 90—100 wm thick. Apothecia up to 1.2 mm
across. Hypothecium 30—45 ym thick. Subhymenium 35 um thick. Hymenium 70 um
high, J+ blue. Asci clavate, with J+ blue tholus (fig. 2), poricidal with extension of the
apex. Ascospores eight, ellipsoid, 10—12 x 6.5—7.5 um. Pycnidia not found. Chemistry:
Atranorin in the upper cortex, lecanoric acid and 2 pigments (as in P. schenckiana
Mull. Arg.) in the medulla.
This lichen is one of a group of very narrow-lobed Parmelias which occur in the
mountains of the southern Cape. At higher altitudes, these mountains are covered with
a good proportion of shrubs with sclerotized, needle-shaped leaves; this type of vegetation
is referred to as fynbos. The Parmelia flora of these mountains seems to have developed
the same way, with a large number of narrow-lobed species present. Even species which
have a wider distribution here produce narrower lobes and a more pulvinate growth
habit at these high altitudes. In the case of the Parmelias, this is almost certainly a re-
sponse to improved mist interception, and is to some extent true for the higher plant
cover as well.
Parmelia fausta resembles P. astricta as well as the newly described P. insignis, but
the most conspicuous difference is the chemistry. P. molybdiza Ny]. is also black below
and contains lecanoric acid, but this is a broader-lobed lichen without any pigments in
the medulla.
At present, this new lichen is known only from the type locality, the Swartberg
range at high altitudes.
186
FIG. 2. Parmelia fausta Brusse, ascus and paraphyses. F. Brusse 4891, holotype. Bar = 10 um.
PARMELIA INSIGNIS Brusse, sp. nov.
Thallus minute, foliosus, saxicola, usque ad 3 cm diametro. Lobi sublineares, 0.1—0.8
mm lati, 100—170 um crassi. Thallus superne cinereus, nitidus, emaculatus, isidiis
sorediisque nullis. Cortex superior 10—17 um crassus. Stratum gonidiale 25-40 um
crassum, algis Trebouxiis, 6—17 um diametris. Medulla alba, 50-90 um crassa. Cortex
inferior 5—20 um crassus. Thallus inferne piceus. Rhizinae simplices, 50—80 um crassae.
Apothecia usque ad 0.8 mm diametris. Hypothecium 15—25 um crassum. Subhymenium
10 wm crassum. Hymenium 60 ym altum, J+ caeruleum. Asci clavati, cum tholis J+
caeruleis. Ascosporae non visae. Pycnidia ignota. Thallus atranorinum et materias
ignotas continens.
TYPUS: SOUTH AFRICA, Cape Province, Robinsons Pass between Mossel Bay and
Oudtshoorn, on TMS outcrops on steep SW slope of Ruitersberg, alt. 790 m, F. Brusse
4801, 4.ii.1986 (PRE, holo-; BM, COLO, LD, MEL, US, iso-), fig. 4.
Thallus, minutely foliose, saxicole, up to 3 cm across. Lobes sublinear, 0.1—0.8 mm
broad, i00—170 um thick. Upper surface grey, nitid, without isidia, soredia or maculae.
187
Upper cortex 10—17 wm thick. Algal layer 25—40 um thick, algae Trebouxia, 6—17 um
diam. Medulla white, 50—90 wm thick. Lower cortex 5—20 um thick. Lower surface
black. Rhizines simple, 50—80 um thick. Apothecia up to 0.8 mm across. Hypothecium
15-25 um thick. Subhymenium 10 ym thick. Hymenium 60 um high, J+ blue. Asci
clavate, tholus J+ blue (only spent asci present). Ascospores not seen. Pycnidia not
found. Chemistry: Atranorin and major amounts of two unidentified substances
(probably meta-depsides), and traces of several others.
Parmelia insignis is another fynbos lichen with sublinear, narrow lobes, and is read-
ily distinguished from its similar looking compatriots by its chemistry. Two major
medullary substances are present which are almost certainly meta-depsides. These sub-
stances could not be positively identified with any of the well known meta-depsides,
but there are a number of recently described meta-depsides, for which no microcrystal
and hydrolysis data, necessary for the rapid confirmation of the identity of these sub-
stances, are available (Chester & Elix, 1978). The substances have consequently been
given code names, and insig.-1 is always the major substance (See Table 1 for R¢ values).
Insig.-2 is present in smaller amounts or absent. Insig.-3 may be a terpene, and is most-
ly present in trace amounts. Up to 6 other traces (best seen in solvent B) occur below
insig.-3, and resemble it in spot colour and shape. This probably represents a series of
closely related terpenes, some of which may be artifacts of the TLC procedure.
At present this lichen is known only from the type locality, the SW slopes of Ruiters-
berg, between Mossel Bay and Oudtshoorn, at the western end of the Outeniqua Moun-
tains.
PARMELIA INUNCTA Brusse, sp. nov.
Thallus minute foliosus, saxicola, usque ad 3 cm diametro. Lobi sublineares, 0.2—0.7
mm lati, 100—120 pm crassi. Thallus superne viridis, nitidus, emaculatus, isidiis sorediis-
que nullis. Cortex superior 15—20 um crassus. Stratum gonidiale 25—35 um crassum, algis
Trebouxiis, 5—16 um diametris. Medulla alba, 35—5O um crassa. Cortex inferior 12—15
um crassus. Thallus inferne piceus. Rhizinae 45—65 um crassae. Apothecia usque ad
0.8 mm diametris. Hymenium non visum. Pycnidia ignota. Thallus acidum usnicum et
acida aliphatica ignota (ut in Parmelia subdecipienti Vain. ex Lynge) continens.
TYPUS: SOUTH AFRICA, Cape Province, Robinsons Pass between Mossel Bay and
Oudtshoorn, on TMS on steep SW slope of Ruitersberg, alt. 790 m, F. Brusse 4810,
4.ii.1986 (PRE, holo-; BM, LD, iso-), fig. 5.
Thallus minutely foliose, saxicolous, up to 3 cm across. Lobes sublinear, 0.2—0.7
mm broad, 100—120 um thick. Upper surface green, nitid, without maculae, isidia or
soredia. Upper cortex 15—20 wm thick. Algal layer 25—35 um thick, algae Trebouxia,
5—16 um diam. Medulla white, 35—5S0 um thick. Lower cortex 12—15 um thick. Lower
surface black. Rhizines 45-65 um thick. Apothecia up to 0.8 mm across. Hymenium
not seen (the available material seemed to have the hymenia grazed off). Pycnidia
unknown. Chemistry: Usnic acid in the upper cortex, and unknown aliphatic acids (as
in P. subdecipiens Vain. ex Lynge) in the medulla.
This is the second small Xanthoparmelia containing the ‘subdecipiens fatty acids’,
the other, Parmelia unctula Brusse (1984), is a more normal Xanthoparmelia with
elongate lobes, and is pale below. These fatty acids do not appear to belong to the con-
stipatic acid series (Chester & Elix, 1979), although I have not personally worked with
188
this group of fatty acids. Brusse (1980) synonomized Parmelia irregularis (Gyeln.)
Kurok. with P. subdecipiens Vain. ex Lynge. P. irregularis, therefore contains the ‘sub-
decipiens fatty acids’. Kurokowa & Filson (1975) depicted a chromatogram on which
the extract of P. irregularis had been run together with those of several other Australian
Parmelias containing the constipatic acid group of fatty acids, and they are shown to
be different. It thus seems certain that the ‘subdecipiens fatty acids’ so common in
southern African Xanthoparmeliae, are not the same as the constipatic acid series, so
common in Australian Xanthoparmeliae.
At present this species is known only from the type collection, from Robinsons Pass
between Mossel Bay and Oudtshoorn.
PARMELIA PUDENS Brusse, sp. nov.
Thallus minute foliosus, saxicola, usque ad 3 cm diametro. Lobi sublineares, 0.1—0.5
mm lati, circa 100 wm crassi. Thallus superne cinereus, nitidus, emaculatus, sorediis
isidiisque nullis. Cortex superior 12—14 um crassus. Stratum gonidiale 15—25 ym cras-
sum, algis Trebouxiis 4.5—17 um diametris. Medulla albida, 40—55 um crassa. Cortex
inferior 10 wm crassus. Thallus inferne piceus. Rhizinae simplices, 40—55 ym crassae.
Apothecia et pycnidia ignota. Thallus atranorinum et acidum salazinicum continens.
TYPUS: SOUTH AFRICA, Cape Province, Robinsons Pass between Mossel Bay and
Oudtshoorn, on TMS on steep SW slope of Ruitersberg, alt. 790 m, F. Brusse 4792,
4.ii.1986 (PRE, holo-), fig. 7.
Thallus minutely foliose, saxicolous, up to 3 cm across. Lobes sublinear, 0.1—0.5 mm
broad, around 100 um thick. Upper surface grey, nitid, without maculae, soredia or
isidia. Upper cortex 12—14 um thick. Algal layer 15—25 wm thick, algae Trebouxia,
4.5—-17 wm diam. Medulla whitish (red in moribund lobes), 40—55 um thick. Lower
cortex 10 um thick. Lower surface black. Rhizines simple, 40—55 um thick. Apothecia
and pycnidia not seen. Chemistry: Atranorin, salazinic acid and a smaller amount of
norstictic acid.
Parmelia pudens resembles the Australian species P. lithophiloides Kurok. (1985) in
character combinations, but the latter has lobes up to 2 mm broad, and is much coarser
than P. pudens, despite being small itself. P. lithophiloides also has constipatic acid in
the medulla while P. pudens is without fatty acids.
This is another fynbos lichen, occurring at higher altitudes in the southern Cape
mountains. At present this species is known only from the type locality, SW slope of
Ruitersberg at the western end of the Outeniqua mountains, between Mossel Bay and
Oudtshoorn.
ACKNOWLEDGEMENTS
The author is grateful to the South African Department of Agriculture and Water
Supply for financial support. Thanks go to Mrs A.J. Romanowski for the photography
and to the typists. Thanks are also extended to Mr. P.W. James for reviewing the manu-
script.
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190
LITERATURE CITED
BRUSSE, F. 1980. A taxonomic and geographic study of the genus Xanthoparmelia in
the Karoo.M.Sc. dissertation, University of the Witwatersrand, Johannesburg, 305 pp.
BRUSSE, F. 1984. New species and combinations in Parmelia (Lichenes) from south-
ern Africa. Bothalia 15 (1 & 2): 315-321.
CHESTER, D.O. & ELIX, J.A. 1978. The identification of four new meta-depsides in
the lichen Ramalina asahinae. Aust. J. Chem., 31: 2745—2749.
CHESTER, D.O. & ELIX, J.A. 1979. Three new aliphatic acids from lichens of genus
Parmelia (subgenus Xanthoparmelia). Aust. J. Chem., 32: 2565-2569.
CULBERSON, C.F. 1972. Improved conditions and new data for the identification of
lichen products by a standardized thin-layer chromatographic method. J. Chromatogr.
72: 113-125.
CULBERSON, C.F. & JOHNSON, A. 1982. Substitution of methyl tert.-butyl ether
for diethyl ether in the standardized thin-layer chromatographic method for lichen
products. J. Chromatogr., 238: 483—487.
HALE, M.E. Jr. 1976. A monograph of the lichen genus Pseudoparmelia Lynge (Par-
meliaceae). Smithson. Contrib. Bot., 31: 1—62.
KUROKAWA, S. 1985. Studies on Australian and Tasmanian species of Parmelia (2).
Bull. natn. Sci. Mus Tokyo, Ser. B (Bot) 11(3): 77—90.
KUROKAWA, S. & FILSON, R.B. 1975. New species of Parmelia from South Austra-
lia. Bull. natn. Sci. Mus. Tokyo, Ser. B (Bot.) 1: 35—48 +4 pl.
Figures 3 to 7.
Fig.3. Parmelia fausta Brusse, habit. F. Brusse 4891, holotype. Scale in mm and cm.
Fig. 4. Parmelia insignis Brusse, habit. F: Brusse 4801, holotype. Scale inmm and cm.
Fig.5. Parmeliainuncta Brusse, habit. F: Brusse 4810, holotype. Scale in mm.
Fig.6. Parmelia eximia Brusse, habit. F. Brusse 4840, holotype. Scale in mm and cm.
Fig. 7. Parmelia pudens Brusse, habit. F. Brusse 4792, holotype. The larger lobed
lichen on the upper margin of this photograph is Parmelia xanthomelaena
Mull. Arg.. Scale in mm and cm.
LST
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4
19
MYCOTAXON
NG Vola nD cee oie 1S October-December 1986
HYPOGEOUS FUNGI FROM THE SOUTHEASTERN UNITED STATES
I. THE GENUS RHIZOPOGON }
Steven L. Miller
Department of Forest Science, Oregon State University, Corvallis, Oregon 97331
SUMMARY
Eleven species of Rhizopogon from the southeastern United States are
redescribed from fresh collections. New information on development and
ectomycorrhizal host range are presented for R. nigrescens, R. fuscorubens, R.
succosus, R. fabri, R. rubescens, R. subaustralis, R. vulgaris, R. couchii, R. evadens ,
R. abietis, and R. truncatus. Rhizopogon atlanticus is redescribed from dried material.
Doubtful species for the southeastern United States are also discussed.
INTRODUCTION
Research on ectomycorrhizae in the southeastern United States has increased
dramatically in the past decade. The need for taxonomic, systematic, and ecological
knowledge of ectomycorrhizal fungi has also increased.
Species in the genus Rhizopogon are hypogeous members of the
Hymenogastrales and are important ectomycorrhizal fungi in the southeastern United
States. Zeller and Dodge (1918) first monographed Rhizopogon in North America, but
no fresh collections were examined from the southeastern United States. In 1928 Coker
and Couch included Rhizopogon in their study of eastern Gasteromycetes. Although
accurate descriptions are provided, this work is no longer a modern treatment, and
several important species are not included. Dr. Alexander Smith in Smith and Zeller
(1966) produced the first modern account of the genus Rhizopogon and included the
major taxa from the southeastern United States. Many of the collections, however, were
described only from herbarium collections. Dr. Smith (Smith and Zeller 1966) admitted
that the greatest disappointment in his investigation of Rhizopogon “was an inability to
satisfactorily identify dried specimens in herbaria" without adequate notes, and he invited
other investigators to apply his concepts to their respective floras.
This paper presents data on Rhizopogon from the southeastern United States.
The ability to study fresh collections in all developmental stages provided me with
insights not previously possible.
1Funded in part by the USDA Forest Service, Pacific Northwest Research Station,
Corvallis, Oregon 97331 U.S.A.
194
METHODS AND MATERIALS
Methods of collection, terminology used, and classification were generally those
of Smith (Smith and Zeller 1966). Colors are from the Methuen Handbook of Colour
(Kornerup and Wanscher 1981) because of its accessibility. Herbarium names are
abbreviated according to Lanjouw and Stafleu (1964). Portions of typical, freshly
collected sporocarps were fixed overnight in 2% glutaraldehyde, dehydrated in ethanol
and embedded in paraffin. After being sectioned, tissues were stained with safranin-fast
green and permanently mounted. Fresh and dried specimens were examined with
standard microchemical reagents, in addition to a saturated aqueous solution of
Ruthenium red. Ruthenium red is a mucilage stain and readily differentiates
brachybasidioles, thick-walled basidia, and certain spore walls. Ultrastructural
investigations and ectomycorrhizal syntheses performed in conjunction with this study
are being published separately.
Key to Rhizopogon species in the southeastern United States
1. Color of fresh, mature peridium dull or bright ago usually overall, but at least over a majority of
the | DEridhw M5 coos seiko cence ccteeVoncss tase cvdedtcdbazedsucsmenscenanuseete ts epseate enc c ine: ateeeae meena 2
1. Color of fresh, mature peridium various, but not distinctly dull or bright yellow overall at any point
IN GEVEIODMENE. 5..--vnsasscelcsncarsacanshconsehecevesssooatensvacensdabentenens seated rededceeeneumsetetstteie tee renee 9
2. Gleba streaked with light-colored veins of sterile tissue; gleba exuding a colorless or pink,
VISCOUS V: LATEX (0. .si se escocesaauantencansesccivevetecenevengseoccestetedensareitte crauhceneenedctteeaeaeens 3
2. Gleba homogenous in color or mottled with lighter or darker colors from insect damage or
erratic maturation, but not streaked with veins of sterile tissue, and not exuding latex
COOSSE SECO OE SSOECS SOOO OE OE OOO ESOS OLOS OHSS OOD OSEO SEES SS OOS OS ODES SOSH SESS ESSE OSES OO OSS OS SOOT OS OSSH ESOS OOSSSOSEOSEESO
3. Fresh peridium light yellow to light orange with abundant, fine rhizomorphs loosely appressed over
the entire surface; not bruising where handled; the gleba dark yellow brown at maturity, not
TANSIUCENE fcc se cescee loll scisnsenootar stu sncnsioewtenvaacuneuccenec suaseateensdwancneeerents 3. R. succosus
3. Fresh peridium greyish yellow to dark greyish yellow, with abundant, fine innate rhizomorphs netting
the surface; frequently flushing red where handled; the gleba brown and nearly translucent at
MALUTILY sorcicvescoccccosensopcdccevecceceaseatpscnwesvadeassecscecdencecssesguessabeoasreemeapesedeereensas 4.R. fabri
4. Spores mostly oblong to oblong-elliptical, thick-walled, with prominent sterigmal remains
1.5-2.0 pm long forming a wide "cupped," truncate base................ccsscssssecsssseees 5
4. Spores mostly subcylindric, subfusoid, ellipsoid or oblong, not noticeably thick-walled,
without prominent sterigmal remains 1.5-2.0 pm long, and usually lacking a
“CUPPEGs UNCALE ’ DASE.......:-00cackescssscsccssssvusetoencsygerteccesemcnseuareeiesntaears eee 6
5. Fresh peridium vivid yellow, even when young; gleba dark yellowish brown at full development;
spores measuring (5.5) 7.0-9.0 (11) x 3.0-5.0 pim...............sccssrccesseseceeeee 11.R. truncatus
5. Fresh peridium white when young, then dull yellow; gleba brownish orange at full development;
spores measuring 7.5-9.0 x 3.0-4.0 pam.uu..........csesccccssscesersrscscesesenesees 12.R. atlanticus
6. Most spores measuring 10.5-13.0 x 3.5-5.0 pumn...............csccessrcesessecsenes 8.R. abietis
6. Most spores measuring less than 10 um long and 2.5-4.0 pm wide, but some spores reaching
10,5=11.0" pi tm: Deng the 22e25o2 15, <5. cassacaasbecasedecedesnsbaceasosesacibaseaneeusevasnsahesaranenecenes 7
7. Rhizomorphs abundant near the base as a tangle of free-hanging, coarse, branching fibrils, and nearly
BDSENC! BDOVE Foidesscskicconssavoccasdecucdavedmatatenntclonosoancaceedeck acecesecculee hive Ean emeee ean eee eee 8
195
7. Rhizomorphs abundant or sparse overall, but not as a tangle of free-hanging fibrils near-the base
PLedanaotn seth awahechavensdits sosvercouede suevuse aroaasnpsesouaNashobenoumeneemt sas ecnaerontetescedosuany 9.R. couchii
8. Spores mostly subcylindric to occasionally subfusoid, measuring 7.5-9.5 (10.5) x 2.5-3.0
pm; with a well-developed turf of inflated cells in the peridium........... 7.R. vulgaris
8. Spores mostly subfusoid and angular, measuring 7.0-10.5 (11) x 3.0-4.0 pm; without a turf
Of inflated cells in ‘the | periditrm) ..5.0......000csccccssecchencnsstvecssdscons 6. R. subaustralis
9. Rhizomorphs abundant to sparse, coarse or fine, conspicuous, turning dark brown or black with
BANGING reo cscs csecese dos tooshoebe civetessscataciewus sass onks das peat ncanacseioes osiloasvbere dane evsaaren ten eivacs 10
9. Rhizomorphs abundant to sparse, but not turning black with handling..................ssccscccssseeereeees 11
10. Fresh peridium tacky; white at full development, easily discoloring where handled to areas
of yellow, pink and orange; rhizomorphs sparse, fine, innate appressed
Pk aatnatees shakuenctas tap eeys oe vadeaeakea Uetae nceccuaidesiesnceesessencenuxdcnseetrs 1.R. nigrescens
10. Fresh peridium dry; greyish orange, brownish orange, or reddish brown at full development,
occasionally bruising red; rhizomorphs abundant, coarse, appressed above, a tangled
network of loosely hanging fibers below..................ccssssesceseres 2.R. fuscorubens
11. Fresh mature peridium pale yellow, bruising red, in age mottled with large areas of brown or dark
brown; spores measuring 7.0-10.5 x(2.5) 3.0-3.5 (4.5) JIM.........sssseseceeseees 5. R. rubescens
11. Fresh mature peridium white, yellowish brown to greyish orange, bruising instantly magenta; spores
measuring (5.5) 6.0-8.0 x 2.5-3.0 (3.5) JIM .........esscccesesseees 10. R. evadens var. evadens
1. Rhizopogon nigrescens Coker and Couch, Gasteromycetes of
Eastern United States and Canada, p. 30. 1928. Figures 1, 11.
ETYMOLOGY: Latin, nigrescens (blackening"), referring to the distinctive
peridium that is white at first, then mottled with a number of colors, and finally drying
blackish or blackish brown.
BASIDIOCARPS: 1.5-4.5(6) cm diam, subglobose, turbinate, compressed or
irregularly lobed. PERIDIUM surface in youth tacky, smooth, tomentose, soon
becoming appressed felty, then unpolished above, easily cracked and wholly removed
from young sporocarps; white when young, then mottled with pale yellow (4A-3),
orange white (5A-2), or pinkish white (7A-2), remaining pale below and in depressions,
becoming brownish orange (6C-5), greyish brown (6D-3) and finally dark brown (7F-6);
bruising intense reddish brown (9D-5) in youth; in cross-section up to 0.5 mm thick, pale
greyish red (9D-5). Rhizomorphs sparse, conspicuous at first, fine, innate-appressed
above in scant reticulum, concolorous with the peridium but brown (6E-6) to dark brown
(6F-6) where handled, inconspicuous as the peridium darkens. Dried peridium black or
dark brown, rhizomorphs inconspicuous. GLEBA: Loculate, locules 1-3 per mm at
full development, smaller when young, irregular in shape, cakelike in consistency in
youth, becoming spongy-cartilaginous in age; white at first, then yellowish grey (3B-2),
or greyish yellow (3B-3), bruising reddish brown (9D-5, 6) where cut in young
specimens. Dried gleba greyish yellow (3B-3 to 4B-3). ODOR sweet, fragrant. KOH
on peridial surface quickly brownish violet (11D-6) to greyish ruby (12D-6); negative to
pinkish white on the gleba. FeSQg on peridial surface slowly dark olive progressing to
black; negative on the gleba.
SPORES: 6.0-9.0 x 2.0-3.0 (3.5) pm, oblong to subfusoid, basal scar present
but not prominent; in KOH hyaline to pale yellowish singly, light olive brown in mass; in
Melzer's reagent hyaline singly, light olive yellow in mass, mostly with 2-3 lipid
droplets. HYMENIAL ELEMENTS: Basidia borne in a distinct hymenium, clavate,
196
15-35 x 5-8 pm, thin-walled and soon collapsing, mostly 4- and 6-spored, 8-spored
common; brachybasidioles 10-30 x 6-12 ym, subglobose, clavate, or obovate, thin-
walled in youth, thick-walled in age, not readily disarticulated in crush mounts.
GLEBA: Trama of hyaline, cylindrical to swollen, loosely interwoven hyphae that are
2-6 (10) ym diam, highly refractive in KOH, thin-walled in youth, mucilaginous with
age; oleiferous hyphae common in mediostratum, 5-15 pm diam, yellowish refractive in
KOH and Melzer's reagent, cylindrical or irregularly swollen and contorted.
Subhymenium cellular, 3-4 cells deep, cells 4-7 pm diam, hyaline, subglobose, elongated
or cubical, mucilaginous with age. PERIDIUM: 370-550 pm thick; of tightly
interwoven, cylindrical, rarely inflated, thin-walled hyphae, 3-10 xm diam, or compacted
into hyphal strands, 10-30 pm diam, 8-15 cells thick, gelatinized in age; heavily
encrusted amorphous pigment that is brownish orange in KOH and Melzer's reagent;
oleiferous hyphae abundant, 5-15 pm diam, yellowish refractive in KOH and Melzer's
reagent, cylindrical or irregularly swollen and contorted. Subcutis lacking. Clamp
connections absent.
HABITAT AND SEASON: Common in the spring and fall from June
through December, associated with 2- and 3-needle pines such as Pinus virginiana, and
P. taeda. Hypogeous or erumpent.
‘COLLECTIONS EXAMINED: LECTOTYPE: North Carolina: Bowlins Creek,
Couch 7582 (NCU). Florida: Alachua Co., W. A. Murrill Autumn 1942, (TENN). Maryland: J.
Trappe 5617, 5618 (OSC). Mississippi: Hancock Co., J. Trappe 4644 (OSC). Virginia:
Montgomery Co., S. L. Miller 695 (OSC).
This species is perhaps the easiest species of Rhizopogon to identify in the
southeastern United States. The small spores, multi-colored mottlings of the fresh
peridium, and the uniform brownish black to black color of the dried peridium are
unmistakable. The fact that the peridium of the younger sporocarps is readily separable
from the gleba is a characteristic that should be compared with R. couchii and R.
evadens also from the southeastern United States, and R. separabilis from the west
coast of the United States.
2. Rhizopogon fuscorubens A.H. Smith, in Smith and Zeller, Mem. N.Y.
Bot. Gard. 14:120-121, 1966. Figures 2, 12.
ETYMOLOGY: Latin, fuscorubens (‘dark brown and reddening"), referring to
the abundant dark brown rhizomorphs, and the reddish brown color often found in the
peridium of older and dried basidiocarps.
BASIDIOCARPS: 1-5 cm diam, subglobose, ovate or turbinate.
PERIDIUM surface dry, smooth, subtomentose in youth, felted below, unpolished
above with age; white at first then pale yellow (4A-3), greyish orange (SB-4), brownish
orange (5C-4, 5 to 6C-6), reddish brown (9C, D-5, 6) or brown (7E-4, 5 to 7F-4, 5) in
age or where exposed; not bruising readily where injured; in cross-section 0.3-0.5 mm
thick, yellowish white (3A-3) to reddish white (7A-2). Rhizomorphs abundant,
circumscribing entire basidiocarp in a reticulate pattern, fine to large, mostly appressed,
some free converging below into a tangled network of loosely hanging strands or single
large rhizomorphic attachment; concolorous at first with fresh peridial surface and
inconspicuous; readily staining brown (7F-4, 5), dark brown (7, 8F-8), or black in age
or where handled, contrasting sharply with the lighter background peridial color. Dried
peridium pale yellow (4A-2, 3) to light brown (7D-5, 6) with abundant brownish red
(8C-5, 6), reddish brown (8D-5, 6), or black mottling; rhizomorphs visible but
conspicuous only on young, pale peridium. GLEBA: Loculate, locules 3-4 per mm at
full development; firm fleshy at first, then spongy-cartilaginous; white in youth, then dull
yellow (3B-3, 4), greyish yellow (4C-4,5), finally light brown (S5D-4,5) with age, often
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mottled with light zones caused by erratic maturation. Dried gleba yellowish white (4A-
2) to greyish yellow (4B-4); hard but easy to section. ODOR absent to faintly metallic.
KOH on peridial surface reddish brown (9F-7, 8); negative on the gleba. FeSOg on
peridial surface negative to slowly grey; negative on the gleba.
SPORES: (6) 7.0-9.5 x 2.5-3.0 pm, oblong to narrowly ellipsoid, basal scar
present but not prominent; in KOH hyaline singly, pale greyish yellow in mass; in
Melzer's reagent hyaline singly, bright orange yellow in mass, mostly with 1-3 false
septa. HYMENIAL ELEMENTS: Basidia borne in a distinct hymenium, narrowly
clavate, 15-20 x 5-8 pm, thin-walled and soon collapsing, mostly 8-spored;
brachybasidioles 10-25 x 6-15 pm, subglobose, clavate or obovate, thin-walled in youth,
moderately thick-walled to thick-walled, mucilaginous in age, not readily disarticulated in
crush mounts. GLEBA: Trama of hyaline, cylindrical to slightly swollen, loosely
interwoven hyphae that are 4-7 (10) pm diam, highly refractive in KOH, thin-walled in
youth, thick-walled, mucilaginous with age; oleiferous hyphae present in mediostratum,
5-12 pm diam, hyaline refractive in KOH, yellowish refractive in Melzer's reagent;
cylindrical or irregularly swollen and contorted. Subhymenium poorly developed,
subcellular at base of some hymenial elements, 1-3 cells deep, cells 6-10 pm diam,
hyaline, subglobose, cubical or elongated, branching, moderately thick-walled with age.
PERIDIUM: 150-250 pm thick. Epicutis of tightly interwoven to appressed hyphae,
3-8 pm diam with hyaline thin to moderately thickened walls; rhizomorphs and hyphal
strands abundant, 15-60 pm diam, 5-15 cells thick, tightly appressed or scattered
throughout, hyphae subparallel in tangential-section, spherical in cross-section,
resembling nests of sphaerocysts; gelatinized in age; heavily encrusted with dark
brownish red amorphous pigment in KOH, pigment readily liquefying into large,
compound reddish brown pigment globules in Melzer's reagent. Subcutis present at the
interface between cuticle and mediostratum of tramal plates and comprising the outer
portion of locules proximal to the cuticle, zone thin, cells mostly anticlinal, 10-15 x 5-8
ym diam, oblong to elongated or swollen, thin-walled, projecting into outermost locules;
not encrusted. Clamp connections absent.
HABITAT AND SEASON: Found late in the fall from October to December
associated with 2- and 3-needle pines such as Pinus serotina and P. rigida. Hypogeous
or erumpent in forests, usually under a deep litter layer.
COLLECTIONS EXAMINED: HOLOTYPE: Oregon: Oswald 10-10-63 (MICH); Lane
Co., J. Trappe 4751, 4836, 4837, (OSC). Massachusetts: Franklin Co., R. E. Halling 3556 (OSC).
Virginia: Princess Anne Co., S. L. Miller 804 (OSC); M. A. Castellano 101 (OSC).
This species is easily recognized by its abundant, conspicuous dark brown to
black rhizomorphs against a pale background peridium color, red to reddish brown
peridium in older or slightly dried specimens and small spores. Dried specimens can
look very similar to R. nigrescens but are generally pale to light yellow or entirely red
mottled with black, rather than entirely black.
3. Rhizopogon succosus A. H. Smith, in Smith and Zeller, Mem. N.Y. Bot.
Gard. 14:115-116, 1966. Figures 3, 13.
=Rhizopogon lowii A. H. Smith in Smith and Zeller, Mem. N.Y. Bot
Gard. 14:55, 1966.
=Rhizopogon superiorensis A. H. Smith, Mich. Bot. 5: 20-21, 1966.
ETYMOLOGY: Latin, succosus ("succulent," "juicy"), referring to the
colorless, viscous latex present in the gleba of mature sporocarps.
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Figs. 3-4. Rhizopogon spp. 3. R. succosus (SLM 723),x 2. 4. R.
fabri (SLM 691), x 2.
BASIDIOCARPS: 1-3 cm diam, mostly subglobose to ovoid or turbinate.
PERIDIUM surface dry, smooth, subtomentose in youth, felted to appressed felty;
pallid white to yellowish white (4A-2) or pale yellow (4A-3) in youth, then light yellow
(4A-4) to light orange (SA-4) in age, mottled with brownish red (9C-6), reddish brown
(9D-6) or brown (6E-6); not bruising where handled; in cross-section, 0.4-0.5 mm thick,
pale yellow. Rhizomorphs abundant over entire surface, fine, loosely appressed above,
free below forming a tangled network of loosely hanging fibrils, often with a single large
rhizomorphic attachment; pale yellow (4A-3), light yellow (4A-4) or light orange (5A-4),
darkening only slightly when handled. Dried peridium uniformly reddish brown (8E-7 to
9E-7). GLEBA: Loculate, locules 3-4 per mm, irregularly shaped; cakelike in
consistency in youth, firm and rubbery in age; exuding a hyaline, viscous latex when cut;
white when young, becoming olive (2E-4), then dark olive brown (4F-8), dark yellowish
brown (5F-4, 5); groups of locules separated by pallid white veins. Dried gleba
unchanged from fresh coloration; glassy hard. ODOR fruity at maturity. KOH on
peridial surface dark brownish red; on the gleba olive green (1E-8). FeSQg on peridial
surface slowly dark olive brown (4E-5); negative on the gleba.
SPORES: 7.5-10.5 (13) x 3.0-4.0 pm, oblong, fusiform-elliptical, naviculate,
reniform or irregularly bent, occasionally subangular or bilobed, smooth, basal scar
prominent; in KOH, pale yellowish olive singly, light olivaceous in mass; in Melzer's
reagent, pale yellowish singly, yellow brown in mass, mostly with one large lipid
droplet. HYMENIAL ELEMENTS: Basidia bore in a hymenial layer or from
various levels within the mediostratum, clavate to irregularly contorted and swollen, 16-
25 x 5-7 pm, thick-walled at first, easily disarticulated in crush mounts, soon collapsing,
1- and 2-spored; brachybasidioles 15-30 x 6-9 (12) ym, clavate to obovate, thin-walled in
youth, thick-walled, mucilaginous in age, easily disarticulated. GLEBA: Trama of
hyaline, cylindrical to swollen and knobby, loosely interwoven hyphae that are 2-5 (8)
pm diam, refractive in KOH and Melzer's reagent, thin-walled in youth, mucilaginous
with age. Subhymenium absent. PERIDIUM: 230-370 pm thick; of tightly
interwoven, cylindrical to inflated, thin- or thick-walled hyphae, 5-15 pm diam; large
highly vesiculose cells common near the surface, 15-30 pm diam, subglobose to
elongated, mostly thick-walled, commonly in small clusters, often tearing to form large
air pockets; hyphal strands abundant, innate appressed or scattered throughout, (15) 20-
30 pm diam, 4-7 cells thick, commonly with one to several large cells surrounded by
smaller cells, hyphae subparallel in tangential section, spherical in cross-section,
resembling nests of sphaerocysts. Subcutis not well differentiated, similar to cutis but
usually lacking large, inflated cells. The entire cutis gelatinized in age, heavily encrusted
with amorphous pigment masses that are magenta to reddish brown in KOH, and
brownish orange in Melzer's reagent. Clamp connections absent.
HABITAT AND SEASON: Common in the fall from October to December,
associated with Pinus strobus, P. taeda, and P. banksiana. Hypogeous under thick
needle-litter layer.
COLLECTIONS EXAMINED: HOLOTYPE: North Carolina: Chapel Hill, Coker
9889 (NCU); 13626 as R. lowii (NCU); Uwharrie Nat. Forest, S. L. Miller 723 (OSC). Michigan:
Luce Co., A. H. Smith 71953 as R. superiorensis (MICH). Tennessee: Blount Co., 9669 (TENN).
Virginia Buckingham Co., S. L. Miller 700 (OSC); Montgomery Co., O. K. Miller 20343 (VPI).
Canada: Ontario, Dunsmore township, C. Godbout 1004 (OSC). Mexico: Chiapas, Lagunas de
Montebello Nat. Park, J. Trappe 3324 as R. superiorensis var. Mexicana (OSC); J. Trappe 3327, 3331
(OSC).
I have collected all developmental stages of this fungus and find that R. succosus ,
R. superiorensis and R. lowii are identical. Rhizopogon lowii corresponds to younger,
less well-developed sporocarps, but the type of R. succosus is overmature and was
probably quite gelatinous when first collected. Of the species synonymized here, R.
superiorensis is the only one that Dr. Smith was able to collect fresh. The color change
from yellow when fresh to brownish red as dried is remarkable for Rhizopogon and
201
admittedly causes confusion in herbarium material. Rhizopogon succosus 1s an
appropriate name because of the viscous latex and the flabby nature of the senescent
gleba.
Two distinct sizes of spores can be found within basidiocarps of R. succosus.
Collections from the southeastern United States, Michigan, and Quebec have a greater
proportion of spores in the larger size and the material labeled R. superiorensis var.
mexicanus Hosford & Trappe has a greater proportion of spores in the smaller size. In
all other respects, the Mexican material agrees with the United States collections and
therefore is recombined as R. succosus A. H. Smith var. mexicanus (Hosford &
Trappe) S. L. Miller (basionym, R. superiorensis var. mexicanus Hosf. & Trappe, Bol.
Soc. Mex. Mic. 14: 11. 1980).
The fresh peridium of R. succosus appears to be mottled with reddish colors but
this is not a staining or bruising reaction. Coloration, in conjunction with the thick-
walled mucilaginous brachybasidioles and glass-hard glebal consistency when dried,
suggests a close affinity to R. luteolus and more proper placement in the Stirps Luteolus
rather than the Stirps Rubescens of Smith's classification.
4. Rhizopogon fabri Trappe, Nova Hedwigia 51:307, 1975. Figures 4, 14.
=Rhizopogon piceus sensu Coker and Couch, Gasteromycetes of Eastern United
States and Canada, p. 34, 1928, non Berkeley andCurtis, Amer. Acad.
Arts Sci. Proc. 4: 129, 1860.
ETYMOLOGY: Latin, genitive of faber (smith), in honor of Prof. Alexander
H. Smith.
BASIDIOCARPS: 1-2 cm diam, mostly subglobose, compressed or reniform,
or occasionally one to several lobed. PERIDIUM surface dry, smooth, subtomentose
in youth, becoming felted, unpolished in age; white to pallid white when young, then
greyish yellow (4B-3), dark greyish yellow (4C-5), remaining white to greyish yellow
below and between lobes, but mottled with brown (6E-7) above; young sporocarps when
handled showing a pale red flush; in cross-section, 0.2-0.3 mm thick, dark greyish
yellow (4C-5) or brown (6E-7). Rhizomorphs abundant, conspicuous in age, fine,
innate fibrils netting the entire surface, 1-3 large, cylindrical rhizomorphs protruding
from basal attachment; concolorous with the young peridium, then dark olive brown (5E-
4) to (SF-8), strikingly darker than peridium. Dried peridium pastel yellow (3A-4)
mottled with large areas of brownish black (7F-4) to black. GLEBA: Loculate, locules
3-4 per mm, filled with spores in age, tough, rubbery; exuding a scant, pinkish yellow,
viscous latex when freshly cut; white at first, then olive (3D-4), finally at maturity brown
(6E-6, 7), nearly translucent; delimited into small groups by whitish or pale yellow veins.
Dried gleba brown (6E-6, 7) to light brown (7E-6, 7); glassy hard. ODOR sweet,
fungal. KOH on peridial surface quickly reddish brown (9D-S, 6); negative on the gleba
or merely darker. FeSOg on peridial surface slowly darker olive than background color;
negative on the gleba.
SPORES: 7.5-11.0 x 3.0-4.0 (4.5) ppm, oblong, fusiform-elliptical, or
irregularly subangular or bent, smooth, basal scar present but not prominent; in KOH,
pale yellow singly, olive brown in mass; in Melzer's reagent, yellowish singly, golden
yellow in mass, mostly with 2-3 lipid droplets; completely filling the chambers in age.
HYMENIAL ELEMENTS: Basidia borne in hyphal stuffed chambers at first, then
from various levels within an irregular hymenium, clavate to elongate-clavate, 15-40 x 5-
7(9) pm, thick-walled at first, soon collapsing, mostly 8-spored, commonly 2-, 4-, and
6-spored; brachybasidioles 15-35 x 7-12 jm, clavate to obovate or subglobose, thin-
walled in youth, thick-walled, mucilaginous in age, easily disarticulated in crush mounts.
GLEBA: Trama of cylindrical to swollen hyphae that are 2-10 pm diam, refractive in
202
KOH and Melzer's reagent, tightly interwoven in the mediostratum, thick-walled, highly
gelatinized near the glebal chambers; oleiferous hyphae common in the mediostratum, 2-5
pm diam, yellowish refractive in KOH and Melzer's reagent, contorted and irregularly
swollen, branching. Subhymenium cellular, not well differentiated, 3-6 cells deep; cells
3-7 ppm diam, hyaline or yellow refractive in KOH and Melzer's reagent, subglobose, to
elongated or irregular, thick-walled, highly gelatinized with age. PERIDIUM: 90-130
thick; of loosely interwoven, cylindrical to inflated, thin- or thick-walled hyphae, 2-8
(10) pm diam near the surface, more tightly compact near the glebal chambers; large,
vesiculose cells common near the surface, thin-walled, readily collapsing to form large air
pockets; oleiferous hyphae common, 2-5 pm diam, yellowish refractive in KOH and
Melzer's reagent, cylindrical to contorted and irregularly swollen, branching; hyphal
strands common, innate-appressed, scattered throughout, 8-20 pm diam, 2-5 cells thick,
generally composed of cells of various sizes, subparallel in tangential-section, spherical in
cross section, resembling nests of sphaerocysts; entire cutis gelatinized in age; heavily
encrusted with amorphous pigment masses that are reddish brown in KOH, yellowish
brown in Melzer's reagent, and readily liquefying into small, yellowish pigment
globules. Subcutis lacking. Clamp connections absent.
HABITAT AND SEASON: Not common, found from summer to late fall,
but mostly November to January, associated with Pinus rigida, and P. palustris.
Hypogeous under deep humus in moist, rich soil.
COLLECTIONS EXAMINED: HOLOTYPE: North Carolina: Chapel Hill, Coker
6069 (NCU); Scotland Co., J. Trappe 8688 (OSC). Virginia: Giles Co., S. L. Miller 519, 691
(OSC); T. M. Flynn 443 (VPI).
This is an unusual fungus. The nearly translucent gleba, viscous latex, and pallid
white veins initially suggest placement in the genus Alpova. In addition, the young
glebal chambers are completely filled with hyphae, at maturity the chambers are
completely filled with spores, and the basidia in older sporocarps arise from various
levels within an irregular hymenopodium. This fungus, however, has a definite layer of
hymenial elements, primarily thick-walled, mucilaginous brachybasidioles, which
preclude its placement in A/pova in the modern sense.
The ultrastructure and development of the locules and hymenium are being
studied to determine the affinities of this fungus. The mucilaginous brachybasidioles and
glass-hard consistency when dried are consistent with Smith's Stirps Luteolus in
Rhizopogon. If this fungus were placed in Alpova it would likely be related to A. klikae
in the subgenus Antridium.
5. Rhizopogon rubescens (Tul.) Tulasne, Giorn. Bot. Ital. 2:58, 1844.
Figures 5, 15.
ETYMOLOGY: Latin, rubescens ("reddening"), referring to the dull red
bruising reaction of the fresh peridium.
BASIDIOCARPS: 1-3 cm diam, subglobose, ovoid, turbinate, or elongate
flattened, furrowed to the base. PERIDIUM surface smooth, dry, cottony fibrillose in
youth, then felted, finally appressed-felted below, areolate above; white when young,
then yellowish white (3A-2) to pale yellow (3A-3, 4A-3), greyish yellow (4B-3) or pale
orange (5A-3), in age mottled with brown (6E-7, 8) or dark brown (6F-7); young
sporocarps bruising greyish red (10B-5) or brownish red (10D-7), older sporocarps
bruising greyish red (9D-5, 6); in cross-section 0.5-1.0 mm thick; greyish red (9D-5).
Rhizomorphs abundant in youth, prominent below and on the sides, innate-appressed,
attached by 1-2 large, cylindrical or flattened rhizomorphs; white or pale yellow (4A-3);
bruising greyish red (9D-5), then dark brown (6F-7). Dried peridium dark brown (6F-8
to 7F-8) or brownish black, with patches or streaks of yellow. GLEBA: Loculate,
203
Figs. 5-6. Rhizopogon spp. 5. R. rubescens (SLM 818), x 2/3. 6.
R.subaustralis (SLM 726), x 1 1/4.
204
locules 1-3 per mm, irregularly shaped, empty at maturity; soft, cakelike at maturity, then
tough, cartilaginous, in age gelatinous, flabby; white at first, then greyish yellow (4C-3,
4) to olive brown (4D-4), often mottled with lighter colors or maturing centrifugally with
lighter colored tissue near the peridium. Dried gleba unchanged from fresh coloration;
waxy, easy to section. ODOR slight, sweet-fungal in youth; in age strong, distinctly of
road tar. KOH on peridial surface reddish brown (8E-7) or (9E-7), in time, dark brown;
negative on the gleba. FeSOg on peridial surface negative, or very slowly olive brown;
negative on the gleba.
SPORES: 7.0-10.5 x (2.5) 3.0-3.5 (4.5) jum, subfusoid, ovate or broadly
ellipsoid, basal scar distinct but not prominent, smooth or minutely punctate; in KOH,
hyaline to yellowish singly and in mass; in Melzer's reagent, hyaline to yellowish singly
and in mass, mostly with 2-3 lipid droplets. HYMENIAL ELEMENTS: Basidia
borne in a distinct hymenium, clavate, 15-35 x 5-7 pm, thick-walled at first, soon
collapsing, mostly 4-spored, 6- and 8-spored basidia also common; brachybasidioles 10-
30 x 6-12 pm, subglobose, clavate, obovate or ellipsoid, thin-walled in youth, thick-
walled, mucilaginous in age, easily disarticulated in crush mounts; basidia,
brachybasidioles, and spores often encrusted or filled with amorphous pigment in locules
near the peridium. GLEBA: Trama of hyaline, highly refractive, cylindrical to slightly
swollen, loosely interwoven hyphae that are 2-8 (10) pm diam, thin-walled in youth,
mucilaginous with age. Subhymenium cellular, 2-4 (5) cells deep, cells 5-8 pm diam,
subglobose to elongated, hyaline, thick-walled with age. PERIDIUM: 240-450 pm
thick. Epicutis of loosely interwoven, cylindrical to slightly inflated, thin or moderately
thick-walled hyphae, 2-5(10) pm diam; hyphal strands frequent, innate-appressed or
scattered throughout, 10-30 pm diam, 2-4 (8) cells thick, hyphae parallel in tangential-
section, spherical in cross-section, resembling nests of sphaerocysts; gelatinized in age,
heavily encrusted with amorphous intra- and intercellular pigment that is reddish brown
to dark brown in KOH, rich reddish brown in Melzer's reagent, and readily liquefies into
small or large pigment globules. Subcutis present at interface between cutis and
mediostratum of tramal plates and comprises the outer portion of locules near the cutis,
zone thin, cells anticlinal, 10-20 x 5-9 ppm diam, oblong, oblong-elliptical or swollen,
mostly thin-walled, projecting into outermost locules; often encrusted with reddish brown
pigment. Clamp connections absent.
HABITAT AND SEASON: Common throughout the fall from October to
December associated with 2- and 3-needle pines such as Pinus virginiana and P. taeda.
Hypogeous, erumpent or fully exposed in soil in well-developed, forested areas.
COLLECTIONS EXAMINED: LECTOTYPE: France, La teste de Buch, Tulasne
October 1843 (K). Michigan: Luce Co. A. H. Smith 58188 (MICH); A. H. Smith 66451 (MICH).
Tennessee: Knox Co., 18122 (TENN); Cades Cove, 20236 (TENN). Virginia: Giles Co., S. L.
Miller 580, 581, 584, 586, 600, 688, 689,690, 815, 818 (OSC); Patrick Co., S. L. Miller 710, 711
(OSC); Roanoke Co., S. L. Miller 579 (OSC). Canada: Quebec, St. Augustine, C. Godbout 104
(OSC); Baie St. Paul, J. Trappe 6364 (OSC).
Confusion surrounding this species will undoubtedly continue until good field
notes are consistently taken on fresh specimens. Traditionally any Rhizopogon species
in the southeastern United States with yellowish coloration and a reddish bruising
reaction has been assigned to R. rubescens. This includes R. subaustralis, R. vulgaris,
R. abietis., and to a lesser extent R. couchii. Rhizopogon rubescens in the sense taken
here has only a pale yellow fresh peridium that is soon mottled with large patches of red
or reddish brown, and becomes areolate in age. Rhizopogon subaustralis, R. vulgaris,
R. abietis, and R. couchii have a pronounced, long-lived yellow stage, with less
tendency to become areolate. In addition, the gleba of R. rubescens and R. couchii
remain quite green (olive) when mature, rather than brown, as in R. subaustralis, R.
abietis, and R. vulgaris.
205
Dried material of R. rubescens and R. subaustralis is difficult to distinguish
without adequate notes because the spores of both species are similar. Rhizopogon
subaustralis spores are generally more fusoid at the apex and are typically more angular
than the mostly oblong or cylindrical spores of R. rubescens . The spores of R. vulgaris
overlap in size with those of R. rubescens but are much narrower. Rhizopogon couchii
spores are shorter than those of R. rubescens, and those of R. abietis are longer and
wider.
6. Rhizopogon subaustralis A. H. Smith in Smith and Zeller, Mem. N.Y.
Bot. Gard. 14:115-116, 1966. Figures 6, 16.
=Rhizopogon brownii A. H. Smith in Smith and Zeller, Mem. N.Y.
Bot. Gard. 14:102-103, 1966.
ETYMOLOGY: Latin, subaustralis ("somewhat southern"), referring to the
distribution limited to the southeastern United States and Mexico.
BASIDIOCARPS: 1-4 cm diam, subglobose, reniform, or elongate flattened,
often irregularly lobed. PERIDIUM surface smooth, dry, cotton-fibrillose in youth,
then felted below, unpolished above in age; white at first, then pale yellow (4A-2),
greyish yellow (4B-4, 4C-3) to pale orange (5A-3), then brownish orange (SC-5-7),
golden brown (SD-5-8), in age brown (7E-4), reddish brown (7E-8) or dark brown (8F-
6) where handled; young sporocarps bruising reddish brown (9D-5, 6); in cross-section
0.5-0.7 mm thick; reddish brown (9D-5). Rhizomorphs not abundant, appressed below,
absent or sparse above, usually attached by one large cylindrical strand; concolorous with
peridium; bruising greyish red (9D-5). Dried peridium brownish orange (SC-4), mottled
with large patches of dark brown (7F-6) to black; rhizomorphs inconspicuous.
GLEBA: Loculate, locules 3-4 per mm, irregularly shaped, empty at maturity; cakelike
in consistency when young, then rubbery, finally cartilaginous; white at first, then pale
yellow (4A-2), and finally greyish yellow (4C-4) or yellowish brown (5E-5, 7). Dried
gleba light brown (SD-4-6) oryellowish brown (SE-5-7). ODOR fungal fruity. KOH
on young peridial surface pale (10A-3) to dull red (10B-4); negative on the gleba.
FeSQq on young peridial surface olive black, often rapidly spreading; negative on the
gleba.
SPORES: 7.0-10.5 (11) x 3.0-4.0 pm, oblong to subfusoid, smooth, basal scar
indistinct, in KOH, hyaline, pale olivaceous singly, in Melzer's reagent, pale yellow
singly, yellow to olive yellow in mass, mostly with 2-3 lipid droplets. HYMENIAL
ELEMENTS: Basidia borne in a distinct hymenium, clavate, 20-35 x 4-6 pm, thin-
walled at first, soon collapsing, 4- and 6-spored; brachybasidioles 5-15 x 5-8 pm,
globose, obovate or ellipsoid, thin-walled in youth, thick-walled, mucilaginous in age,
easily disarticulated in crush mounts. GLEBA: Trama of hyaline, loosely interwoven
hyphae that are 3-9 (12) pm diam, thin-walled in youth, mucilaginous with age;
oleiferous hyphae common in the mediostratum and near the peridium, 5-20 pm diam,
yellowish refractive in KOH and Melzer's reagent, cylindrical to convoluted, branched.
Subhymenium cellular, 2-3(4) cells deep, cells 3-7 pm diam, hyaline, subglobose or
elongated. PERIDIUM: 370-500 pm thick; of loosely interwoven, cylindrical to
slightly inflated, thin- or moderately thick-walled hyphae, 2-5 pm diam; hyphal strands
common, innate-appressed or scattered throughout, 3-10 (15) pm diam, 3-5 cells thick,
subparallel in tangential-section, spherical in cross-section, resembling nests of
sphaerocysts; oleiferous hyphae common, 3-20 ym diam, yellowish refractive in KOH
and Melzer's reagent, cylindrical or contorted, branched; cutis gelatinized in age, heavily
encrusted with amorphous pigment that is reddish brown in KOH, olive brown in
Melzer's reagent, readily liquefying into small or large pigment globules. Subcutis
lacking. Clamp connections absent.
206
HABITAT AND SEASON: Occurring spring and summer, very common in
the fall from October to November associated with 2- and 3-needle pines such as Pinus
virginiana, P. rigida, and P. echinata. Hypogeous, erumpent, or fully exposed, usually
in early successional, poorly developed forests, on rocky slopes, or disturbed areas.
COLLECTIONS EXAMINED: HOLOTYPE: North Carolina: Chapel Hill, 7450
(NCU); Uwharrie Nat. Forest, §. L. Miller 724, 726 (OSC). Maryland: J. Trappe 5616 (OSC).
Louisiana: Livingston Parish, S. M. Zeller 7203 as R. brownii. Mississippi: Harrison Co., J.
Trappe 4642 (OSC). Virginia: Montgomery Co., S. L. Miller 717, 718, 719 (OSC).
The importance of this species as an ectomycorrhizal fungus can only be assumed
from the great abundance of its basidiocarps in the fall. Rhizopogon subaustralis differs
from many species of Rhizopogon; it consistently occurs on disturbed or eroded banks
and in early successional reforestation sites. Rhizopogon subaustralis would be a good
candidate for inoculation onto nursery stock intended for less than optimal sites.
7. Rhizopogon vulgaris (Vitt.) M. Lange, Dansk Bot. Ark. Bd. 16, nr. 1,
p. 56, 1956. Figures 7, 17.
ETYMOLOGY: Latin, vulgaris ("“common," “ordinary"), referring to the fact
that this fungus is widely distributed and commonly collected worldwide.
BASIDIOCARPS: 1-4 cm diam, subglobose, ellipsoid, flattened or irregularly
lobed, frequently compound. PERIDIUM surface dry, smooth, cottony fibrillose in
youth, then felted, shiny, metallic above in age after prolonged exposure; white when
young and unexposed, then yellowish white (3,4A-2), pastel yellow (3A-4), finally
greyish yellow (4C-S, 6) to olive brown (4D-5,6); white peridium bruising reddish white
(9A-2), older or exposed peridium bruising brownish red (10D-6, 7) to violet brown
(11E-5, 6); in cross-section 0.5-1.5 mm thick, thickest near the basal attachment,
flushing dull red (10B-4). Rhizomorphs abundant near basal attachment, coarse, free
hanging, profusely branched, rarely appressed over upper surface; white bruising reddish
white (9A-2) when young, in age white to yellowish white (4A-2), often with no
bruising reaction. Dried peridium greyish yellow (4B-3; 4C-4), often with patches of
brighter yellow; rhizomorphs yellowish brown (5F-8 ), appressed, conspicuous near the
base. GLEBA: Loculate, locules 4-5 per mm at full development; soft, cakelike at first,
then tough, cartilaginous, in age gelatinous; white to yellowish white (2A-2) or pale
yellow (2A-3) in youth, then olive yellow (3C-6), finally olive brown (4D, E-7) to
yellowish brown (SE-4-8), often mottled light and dark because of erratic maturation;
young specimens with pale gleba occasionally bruising pinkish white (7A-2). Dried
gleba greyish yellow (4B-4) to yellowish brown (SE-8). ODOR similar to the
commercial mushroom fresh, with age strongly of road tar. KOH on white peridium
reddish white (9A-2), on yellow or olive areas brown (7E-7,8); negative on the gleba.
FeSOg on white peridium dull grey to blackish grey, negative on older peridium;
negative on the gleba.
SPORES: 7.5-9.5 (10.5) x 2.5-3.0 pm, subcylindric, oblong to narrowly
subfusoid, often curving slightly in profile, basal scar present but not prominent, in
KOH, pale yellowish singly, dull yellowish grey in mass; in Melzer's reagent, pale
yellow singly, dull olive yellow in mass, mostly with 2-3 lipid droplets. HYMENIAL
ELEMENTS: Basidia borne in a distinct hymenium, subcylindric to narrowly clavate,
12-18 x 4-5 jim, thin-walled and soon collapsing, mostly 8-spored; brachybasidioles 10-
20 x 6-10 pm, subglobose, clavate or obovate, thin-walled in youth, thick-walled,
mucilaginous in age, not readily disarticulated in crush mounts. GLEBA: Trama of
hyaline, cylindrical to slightly swollen hyphae that are 4-7 pm diam, highly refractive in
KOH, thin-walled in youth, mucilaginous with age; oleiferous hyphae present in
mediostratum, 6-12 jim diam, hyaline refractive to deep yellow in KOH and Melzer's
207
wan ys
4
‘\
rea
\ Ae \
a
tigs. 7-8. Rhizopogon spp. 7. R. vulgaris (SLM 860), x 2/3. 8. R.
‘ouchii (SLM 716), x 4/5.
208
reagent, cylindrical or irregularly swollen and contorted. Subhymenium poorly
developed, composed of hyaline, thin- or thick-walled, branching, cylindrical or cubical
hyphae. PERIDIUM: 300-360 pm thick. Epicutis a turf of globose to ovoid, thin-
walled, inflated cells. Subcutis a layer of appressed, interwoven, thin-walled cylindrical
hyphae, 6-14 pm diam; oleiferous hyphae abundant, 8-20 pm diam, dark yellowish
brown in KOH and Melzer's reagent, cylindrical or irregularly swollen and contorted,
occasionally branching; slightly gelatinized in age, lightly encrusted with amorphous
pigment that is orange brown to reddish brown in KOH, orange brown in Melzer's
reagent, readily liquefyinginto large, compound orange brown pigment globules . Clamp
connections absent.
HABITAT AND SEASON: Common in late summer to fall from July to
December associated with a variety of conifers including Pinus virginiana, P palustris,
and P. elliotii in the southeastern United States, and P. ponderosa, P. contorta, P.
monticola, Tsuga mertensiana, and Abies amabilis in the western United States.
Hypogeous to erumpent, usually in disturbed, harsh forest sites or eroded slopes.
COLLECTIONS EXAMINED: Arkansas: Montgomery Co., J. Trappe 6660, 6661 (OSC);
Union Co., J. Trappe 7392, 7393 (OSC). California: Siskiyou Co., S. L. Miller 860 (OSC); J.
Trappe 4020, 5639 (OSC). Florida: Tallahassee, 21068 (TENN). Massachusetts: Franklin Co.,
R. E. Halling 3555 (OSC). Mississippi: Picayune, W. Cibula 648, 650 (OSC). Oregon:
Deschutes Co., J. Trappe 4503, 7666 (OSC); Union Co., J. Trappe 4757 (OSC). Tennessee: Knox
Co., 6609 (TENN). Virginia: Montgomery Co., S. L. Miller 587 (OSC).
Rhizopogon vulgaris is similar to R. subaustralis, occurring primarily on
disturbed, eroded, or less than optimal sites. Often even large basidiocarps when cut
open will be immature, and mature basidiocarps are often connected via tangled
rhizomorphs to dozens of small, white, flattened or irregularly shaped primordia.
As with Rhizopogon succosus and other species of Rhizopogon, the spores of
R. vulgaris exhibit different spore-length ranges in different regions of North America
and Europe. Svrcek (1958) describes several varieties of R. vulgaris to account for
variation in spore length. Rhizopogon vulgaris var. intermedius Svrcek best fits the
mean spore length of most of the material that I have examined from the southeastern
United States. The type collection of R. vulgaris var. intermedius cited by Svrcek,
however, is mentioned under R. roseolus var. intermedius in the front of the book, and
validly described in Latin under R. vulgaris var. intermedius in the keys at the back of
the book. I am not certain of Svrcek's intent, and because spore length in the
southeastern United States material is highly variable and overlaps with several of the
varieties described by him, I am not ascribing varietal status.
8. Rhizopogon abietis A.H. Smith, in Smith and Zeller, Mem. N.Y. Bot.
Gard. 14:93-94, 1966. Figure 18.
ETYMOLOGY: Latin, abietis (‘'fir-associated”), referring to the collection of the
type specimen of this species close to members of the conifer genus Abies.
BASIDIOCARPS: 1.5-4 cm diam, subglobose, ovoid or pyriform.
PERIDIUM surface dry, to slightly tacky in aging sporocarps, smooth, cottony
fibrillose in youth, then appressed felty; white when young or unexposed, pale yellow
(4A-3) to light yellow (4A-4) or reddish yellow (4B-6) in age or where exposed; bruising
pale red (9A-2) at first, bruised areas quickly turning brown (6E-8 to 7E-4) or reddish
brown (8E-5-7); in cross-section 0.3-0.5 mm thick, bruising greyish red (7B-4).
Rhizomorphs abundant below, appressed, converging into one or two coarse
rhizomorphs at point of attachment, scattered to absent above; concolorous with
peridium; readily bruising dull red (8B-3). Dried peridium light yellow (4A-4) to brown
209
(6E-6), often with patches of black. GLEBA: Loculate, locules 4-5 per mm at full
development; cartilaginous even in youth, gelatinizing in age; white at first, then greyish
yellow (4C-5), yellowish brown (5D-S to 5F-4, 5, 6) in age. Dried gleba light brown
(5C,D-5, 6) to brown (6E-5, 6); hard to bone hard but easy to section. ODOR faint,
fungal-fruity when young, distinctly of road tar in age. KOH on peridial surface reddish
brown (8E-7); negative on the gleba. FeSOq on the peridium slowly greyish olive (4E-
3); negative on the gleba.
SPORES: 10.5-13 (15.5) x 3.5-5.0 (5.5) pm, mostly subfusoid, ranging to
ellipsoid or ovoid, often slightly curving in profile view, basal scar prominent, fine
remnants of the sterigma projecting slightly beyond a truncate base; narrowly clavate
spores, possibly conidia also present, 15-22 x 3-3.5 jim, truncate at base but mostly
lacking sterigmal projections; in KOH, hyaline singly, pale olive grey in mass; in
Melzer's reagent, pale yellow singly, greyish olive in mass, mostly with 2 lipid droplets.
HYMENIAL ELEMENTS: Basidia borne in a distinct hymenium, 15-20 x 5-7 pm,
mostly thin-walled and soon collapsing, but many with moderately thick walls supporting
conidial spores, 2-, 4-, and 6-spored; brachybasidioles 8-20 x 6-10 pm, subglobose,
clavate, or obovate, thin-walled in youth, thick-walled, mucilaginous in age, often with
amorphous yellow refractive contents in KOH and Melzer's reagent, readily disarticulated
in crush mounts. GLEBA: Trama of hyaline hyphae that are 4-7 pm diam, moderately
refractive in KOH, cylindrical to swollen near septa, loosely interwoven, thin-walled in
youth and in age, gelatinous; oleiferous hyphae abundant in mediostratum, 3-8 pm diam,
hyaline refractive in KOH, yellowish refractive in Melzer's reagent, cylindrical or
irregularly swollen and contorted, often branching. Subhymenium subcellular but poorly
developed, composed of cubical or cylindrical and branching hyphae, thin-walled at first,
thick-walled, mucilaginous in age. PERIDIUM: 220-260 pm thick; of hyaline, thin-
walled, loosely interwoven, cylindrical hyphae, 4-10 jm diam, large inflated cells up to
15 pm diam common, with a turf of clavate terminal cells, 25-40 x 10-14 jm, cuticle not
readily gelatinizing in age; interior portion lightly encrusted with yellow to orange yellow
amorphous pigment in KOH and Melzer's reagent, turf cells free from encrusting
material. Subcutis absent. Clamp connections absent.
HABITAT AND SEASON: Found in the fall from September to December.
This species is associated with 2- and 3-needle pines such as Pinus virginiana and P.
taeda in the southeastern United States, but is reported with Tsuga, Abies and Picea
from the western states. Hypogeous under deep litter in well-developed forest cover.
COLLECTIONS EXAMINED: HOLOTYPE: Idaho: Valley Co., A. H. Smith 65348
(MICH). California: Siskiyou Co., J. Trappe 5374, 7603 (OSC). Oregon: Lane Co., J. Trappe
4654 (OSC); Marion Co., J. Trappe 876 (OSC). Pennsylvania: Clarion Co., J. Trappe 7717 (OSC).
Tennessee: Knoxville, 19494 (TENN); Hardin Valley, 21614 (TENN). Virginia: Montgomery Co.,
S. L. Miller 589 (OSC).
Rhizopogon abietis is most similar to fresh collections of R. vulgaris in that both
species are distinctly yellow, bruise red, and have a tangle of white rhizomorphs near the
point of attachment. In R. abietis , however, the much longer, wider basidiospores,
frequently with a knobby, crooked apex, are distinctive. The exceedingly long, thin
spores found among the basidiospores are also distinctive and may be conidial in origin.
9. Rhizopogon couchii A. H. Smith, in Smith and Zeller, Mem. N.Y. Bot.
Gard. 14:115-116, 1966. Figures 8, 19.
ETYMOLOGY: In honor of Dr. John N. Couch, who has collected hypogeous
fungi in the southeastern United States for many years, and who refers to himself as
William C. Coker's "truffle pig."
BASIDIOCARPS: 1-6 cm diam, subglobose, turbinate, angular, irregularly
elongate and flattened, or lobed. PERIDIUM surface dry, smooth, subtomentose to
felty in youth, then unpolished above, easily separable from gleba; white or yellowish
white (4A-2) when young or covered with soil, then pale yellow (4A-3), light yellow
(4A-4), or greyish yellow (4B-4, 5); in age or where handled, brownish orange (5C-4;
6C-3) or brown (6E-6), pale yellow (4A-3) below and in depressions; young sporocarps
bruising reddish brown (9D-4), older sporocarps bruising light brown (7D-6), brown
(7D-7) or reddish brown (8D-5); in cross-section 0.3-1.0 mm thick, reddish brown (9D-
5). Rhizomorphs abundant, large, appressed below, fine, more reticulate above, fused to
a single point of attachment; concolorous with peridium. Dried peridium brown (7E-7)
or dark brown (7F-8); rhizomorphs inconspicuous. GLEBA: Loculate, locules 4-5 per
mm in youth, in age larger, 2-3 per mm, irregularly shaped, empty at maturity; cakelike
in consistency when young, then firm, moist, finally dry, cartilaginous; white at first,
then dull yellow (3B-3), greyish yellow (3B-5 to 3C-3), finally olive brown (4D-5),
maturing centrifugally; often bruising pale red (9A-3) to greyish red (9C-5) where
exposed. ODOR fruity. KOH on peridial surface deep red to reddish brown; on gleba
greyish red (8B-4, 5). FeSOq on peridial surface slowly yellowish brown (S5E-5-7 to
S5F-5-7) or olive black; negative on the gleba.
SPORES: 5.5-8.5 x 2.0-3.0 pm, oblong to oblong-elliptical, smooth, basal scar
indistinct; in KOH, hyaline singly, yellowish to olivaceous in mass; in Melzer's reagent,
yellowish singly, olive yellow in mass, mostly with 2 lipid droplets. HYMENIAL
ELEMENTS: Basidia borne in a well-developed hymenial layer, mostly clavate, 10-30
x 4-7 pm, thin-walled at first, soon collapsing, 4-, 6- or mostly 8-spored;
brachybasidioles 6-10 x 5-6 pm, subglobose to obovate, hyaline, thin-walled in youth,
thick-walled in age, gelatinized and pigment-filled or encrusted in age in locules proximal
to the peridium. GLEBA: Trama of hyaline, cylindrical to inflated hyphae that are 2-5
pm diam, subparallel to interwoven, or pseudoparenchymatic adjacent to the chambers,
thin-walled in youth, mucilaginous in age. Subhymenium cellular, 2-3 (4) cells deep,
cells 4-10 pm diam, pale orange or yellow orange in KOH, yellowish in Melzer's
reagent, globose, subglobose, or elongate. PERIDIUM 280-300 pm thick; of tightly
interwoven, cylindrical or slightly inflated, thin-walled hyphae, 4-10 (15) pm diam;
enlarged cells scattered near glebal chambers; hyphal strands present, innate-appressed or
submerged, 20-40 jm diam, 3-10 cells thick, parallel in tangential-section, spherical in
cross-section, resembling nests or sphaerocysts; gelatinized in age, heavily encrusted
with amorphous pigment that is yellow to yellow brown in KOH, intense yellow to
yellow brown in Melzer's reagent, intra- and intercellular near the surface, intercellular
only near the locules. Subcutis lacking. Clamp connections absent.
HABITAT AND SEASON: Common Iate in the fall from October to
December associated with 2- and 3-needle pines such as Pinus virginiana, P. taeda, and
P. resinosa. Hypogeous or erumpent in forests, usually under a deep litter layer.
COLLECTIONS EXAMINED: HOLOTYPE: North Carolina: Chapel Hill, 6003
(NCU). Florida: Tallahassee, 20273 (TENN). Tennessee: Cades Cove, 4860 (TENN). Virginia:
Montgomery Co., S. L. Miller 715, 716 (OSC). West Virginia: Pocahontas Co., S. L. Miller 504,
554 (OSC); O. K. Miller 19566 (VPI).
Rhizopogon couchii varies considerably in color. I have seen large, fully mature
basidiocarps that are white, yellow, or brown, depending on the depth at which the
sporocarps were found. The more exposed the peridium is, the darker the color. The
abundant appressed rhizomorphs and yellowish color somewhere on the fresh peridium
are enough to distinguish this species, however. The red bruising of the gleba is best
demonstrated in fresh, moist, undamaged sporocarps.
Rhizopogon couchii, R. vulgaris, and R. evadens are similar when dried and are
best identified when fresh. Rhizopogon. couchii has abundant rhizomorphs overall, but
A ag
not white, loosely hanging rhizomorphs as in R. vulgaris. Rhizopogon evadens has
only sparse rhizomorphs, is typically white, pale yellow to greyish orange and has a
silky, greyish red flush overall. Rhizopogon vulgaris has a turf of inflated, ovoid
hyphal end-cells in the peridium.
Rhizopogon smithii Hosford from coastal regions of the Pacific northwest shows
similar color variation, rhizomorphic pattern, and red bruising of the gleba as in R.
couchii. In addition, the spores of R. smithii are the same size as those of R. couchii.
Further examination of these two species may warrant synonymy.
10. Rhizopogon evadens A.H. Smith var. evadens in Smith and
Zeller, Mem. N.Y. Bot. Gard. 14:151-152, 1966. Figures 9, 20.
ETYMOLOGY: Latin, evadens (‘escaping,” "going out"), possibly because
basidiocarps of this fungus are frequently erumpent.
BASIDIOCARPS: 1-5 cm diam, subglobose to elongate flattened, frequently
irregularly lobed and folded. PERIDIUM surface dry, smooth, subtomentose at first,
then appressed felty, often scurfy, in age silky; easily separable from gleba; white to
yellowish white (4A-2) when young or unexposed, often pale yellow (4A-3) to yellowish
brown (5D-S, 6) in patches, finally greyish orange (SB-3, 4 to 6B-3, 4); white peridium
bruising greyish Magenta (13C, D-5, 6), older or exposed peridium discoloring to
greyish red (7B-3, 4) or reddish brown (8 9D-5) where handled; in cross-section 0.5 to
1.0 mm thick, thickest near the base, white, flushing immediately greyish red (9C-4, 5).
Rhizomorphs abundant, appressed, coalescing into a single large rhizomorphic
attachment below, tapering to fine, scant fibrils above; concolorous with peridium at first,
then brown (7E-5, 6) where handled. Dried peridium pale orange (5A-3) to light orange
(SA-4) below and in depressions, brown (6E-4, 5 or 7E-5, 6) to dark brown (6F-4, 5)
elsewhere; rhizomorphs dark brown (6F-4) and inconspicuous. GLEBA: Loculate,
locules 4-5 per mm when young, up to 2-3 per mm at full development; soft, cakelike
young, then tough, cartilaginous in age; white in youth, then greyish yellow (2C-3; 3B-5
to 3C-4), maturing centripetally with pale coloration proximal to the peridium; young or
mature gleba often bruising pale red (9A-3) to dull red (9C-4). Dried gleba pale yellow
(4A-3) to light yellow (4A-4), hard but easy to section. ODOR not distinctive to acidic
disagreeable in age. KOH on peridium reddish brown (9D-6, 7); on young gleba often
pinkish white (9A-2). FeSO4 on peridium slowly olive; negative on the gleba.
SPORES: (5.5) 6.0-8.0 x 2.5-3.0 (3.5) pm, narrowly oblong to ovoid,
frequently curved in profile, basal scar present but not prominent; in KOH, hyaline to
pale yellow singly, pale yellowish grey in mass; in Melzer's reagent, pale yellow singly,
olive yellow in mass, mostly with 2-3 lipid droplets. HYMENIAL ELEMENTS:
Basidia bore in a distinct hymenium, 15-25 x 5-6 pm, thin-walled and soon collapsing,
narrowly clavate, 4-, 6-, or 8-spored; brachybasidioles 8-25 x 6-15 pm, subglobose,
clavate or obovate, thin-walled to moderately thick-walled in youth, thick-walled,
mucilaginous in age, readily disarticulated in crush mounts. GLEBA: Trama of
hyaline, cylindrical to slightly swollen, loosely interwoven hyphae that are 3-8 jm diam,
refractive in KOH, thin-walled in youth, highly gelatinous in age; oleiferous hyphae
common in mediostratum, 4-13 pm diam, hyaline refractive in KOH, yellow refractive in
Melzer's reagent, cylindrical or irregularly swollen and contorted, branching.
Subhymenium poorly developed, composed of hyaline, thin-walled, cubical, elongated
or branching cylindrical hyphae. PERIDIUM: 200-320 pm thick; epicutis of thin-
walled, tightly appressed, parallel, gelatinous hyphae, 3-5 pm diam, heavily encrusted
with dull yellow to orange yellow amorphous pigment in KOH, bright yellow orange in
Melzer's reagent. Subcutis present but not well differentiated from epicutis, of thin-
walled, cylindrical, non-gelatinous hyphae, 5-25 sm diam, with abundant hyphal strands
appearing in cross-section as nests of sphaerocysts; oleiferous hyphae abundant, 5-12 pm
ps Ws
diam, hyaline refractive in KOH, yellow refractive in Melzer's reagent, cylindrical or
irregularly swollen and contorted, branching, lightly encrusted with dull yellow pigment
or free from encrusting material. Clamp connections absent.
HABITAT AND SEASON: Found in the spring, summer and fall, but
mostly from October to December. This species is associated with 2- and 3-needle pines
such as Pinus virginiana, P. taeda, P. palustris, and P. elliotii in the southeastern United
States, and P. ponderosa, P. contorta, P. lambertiana, and Pseudotsuga menziesii in
the west.
COLLECTIONS EXAMINED: HOLOTYPE: Idaho: Custer Co., A. H. Smith 65484
(MICH). Colorado: Boulder Co., J. Trappe 5700, 8055 (OSC). Mississippi: Pearl River Co., W.
Cibula 648, 649, 650, 656, 666, 669 (OSC); Santa Rosa, W. Cibula 670 (OSC); Saucier, W. Cibula
677 (OSC). North Carolina: Uwharrie Nat. Forest, S. L. Miller 725 (OSC). Ohio: Hocking Co.,
W. B. Cooke 41669 (OSC). Oregon: Deschutes Co., J. Trappe 8185 (OSC); Douglas Co., J. Trappe
343 (OSC); Jackson Co., J. Trappe 6285 (OSC); Josephine Co., J. Trappe 7040, 7429 (OSC).
Tennessee: Cades Cove, 18150 (TENN). Virginia: Montgomery Co., S. L. Miller 705 (OSC).
Rhizopogon evadens is most often confused with R. couchii or R. vulgaris
because of the small spores. Rhizopogon evadens , however, lacks the distinctly yellow
peridium characteristic of the other two species in the field and typically will bruise much
faster and turns more Magenta when young and fresh.
11. Rhizopogon truncatus Linder, Rhodora 26:196-197, 1924.
Figures 10, 21.
=Rhizopogon cokeri A. H. Smith in Smith and Zeller, Mem. N.Y. Bot
Gard. 14:58-59, 1966.
ETYMOLOGY: Latin, truncatus ("cut off"), referring to the truncate,
prominent basal scar on the spores.
BASIDIOCARPS: 0.5-3 cm diam, globose, subglobose, or ovoid.
PERIDIUM surface dry, granulose to pulverulent, unpolished where handled; yellow
(3A-6) to vivid yellow (3A-8) even in youth, greyish yellow (2B-7 to 3B-7) or olive
yellow (3C-7) where handled; not bruising; in cross-section 0.3-0.5 mm thick, yellow
(3A-6). Rhizomorphs sparse, inconspicuous, fine, reticulating above, 1-3 large strands
appressed below, concolorous with peridial surface. Dried peridium yellow (3A-6,7),
often mottled with olive yellow (3C-7) or deep orange (6A-8). GLEBA: Loculate;
locules small, 4-6 per mm, irregularly shaped, empty at maturity; soft, cakelike in
consistency when young, firm in age; yellowish white (4A-2) to pale yellow (4A-3) at
first, then greyish yellow (4B-4), dark olive brown (4F-8) or yellowish brown (SE-8), .
finally dark yellowish brown (6F-8). Dried gleba unchanged in color; easy to section.
ODOR slightly oily or not distinctive. KOH negative on the peridial and glebal surface.
FeSOgq negative on peridial and glebal surface.
SPORES: (5.5) 7.0-9.0 (11) x 3.0-5.0 pm, oblong, oblong-elliptical,
cylindrical, or ellipsoid, frequently irregularly allantoid, reniform, subangular or bilobed,
smooth truncate, the basal scar and sterigmal remains prominent, walls thickened at
maturity; in KOH, hyaline to pale yellow singly, yellowish brown in mass; in Melzer's
reagent, immature spores hyaline to strongly dextrinoid singly and in mass, mature
spores pale yellowish brown to slightly dextrinoid in mass; mostly with 1-2 lipid
droplets. HYMENIAL ELEMENTS: Basidia borne in hymenial layer, clavate, 11-14
x 4.5-5.5 pm, hyaline, thick-walled at first, then autolyzing, 6- or 8-spored;
brachybasidioles subglobose to obovate, 7-10 x 6-8 pm, thick-walled, gelatinizing at
maturity. GLEBA: Trama of hyaline, cylindrical to swollen hyphae that are 2-5 pm
21S
Figs. 9-10. Rhizopogon spp. 9. R. evadens var. evadens (SLM
705), x 4/5. 10. R. truncatus (Dan Luoma 638), x 1 1/4.
214
diam, thin-walled in youth, mucilaginous, refractive in age, subparallel in the
mediostratum, interwoven and gelatinized adjacent to the hymenium; scattered oleiferous
hyphae present in the mediostratum, 5-10 (15) pm diam, refractive or yellowish refractive
in KOH and Melzer's reagent. PERIDIUM 100-220 pm thick; of uniformly cylindrical
to occasionally swollen , branching, anastomosing, thin-walled hyphae, 4-7 pm diam;
end cells 3-40 x 4-6 ppm diam, blunted or rounded at the tip, protruding or intertwined; in
KOH, pale yellow, the pigment diffusing into the fluid; bright yellow in Melzer's
reagent, the pigment lightly encrusting the hyphae, not diffusing into the fluid. Subcutis
lacking. Clamp connections absent; short, stubby branches resembling false-clamps
common.
HABITAT AND SEASON: Fairly common throughout the summer from
July to November and occasionally in the spring. In the southeastern United States, this
species is most commonly reported from forests containing Tsuga canadensis, but it also
occurs with Pinus resinosa. On the west coast it is associated with Tsuga mertensiana,
Pseudotsuga menziesii, and Abies spp. Hypogeous or imbedded in well-rotted conifer
logs, or woody debris.
COLLECTIONS EXAMINED: HOLOTYPE: New Hampshire: D. H. Linder July
1922 (FH). California: Del Norte Co., J. Trappe 3000 (OSC); Siskiyou Co., J. Trappe 4588 (OSC).
North Carolina: Highlands, Coker 10511, as R. cokeri (NCU); Blount Co., V. Cotter 745 (VPI).
Oregon: Douglas Co., J. Trappe 350, 5945 (OSC); Jackson Co., J. Trappe 2893, 4694 (OSC);
Josephine Co., J. Trappe 7051(OSC); Klamath Co., J. Trappe 5264 (OSC); Lane Co., J. Trappe 2183,
4652, 5933 (OSC). Canada: Nova Scotia, K. A. Harrison 7670 (MICH).
Rhizopogon truncatus is interesting and unusual for several reasons. Unlike
most Rhizopogon species the pigment in the peridium is bright yellow, pulverulent and
dissolves readily in ETOH, KOH, and only slightly less readily in water. The spores are
hyaline to pale yellow when young, then strongly dextrinoid, and finally pale yellow
brown to only slightly dextrinoid when older. The gleba reaches full development slowly
and in most collections the color is pale yellow. At full development, however, the gleba
is dark yellowish brown.
The type of R. truncatus was collected by Linder (1924) from the top of a stump
among hemlock trees in a birch-hemlock woods, where it had presumably been left by a
small mammal. The length of time it had been there is unknown.
Unpublished data from the author shows that R. truncatus spores have 4-6 wall
layers, several of which develop after the spore is released from the basidium. The
spores of Linder's original material are mostly collapsed and hyaline to pale yellow, as if
they are young. The types of R. truncatus and R. cokeri differ only developmentally,
and neither are fully mature. The type collection of R. truncatus is the younger of the
two and perhaps weather-worn from its exposure on the stump. The difference in
coloration between the two dried collections is conceivably caused by rainfall washing a
portion of the bright yellow pigment from Linder's collection.
12. Rhizopogon atlanticus Coker and Dodge in Coker and Couch,
Gasteromycetes of Eastern United States and Canada, p. 35, 1928. Figure 22.
ETYMOLOGY: Latin, atlanticus ("near the Atlantic") referring to the
distribution limited to the southeastern United States bordering the Atlantic Ocean.
BASIDIOCARPS: 1-4 cm diam, subglobose, elongate, or ovoid. Peridium
surface white when young, dull yellow with age; not bruising where handled.
Rhizomorphs present but not abundant, large, innate-appressed, a single strand at point
of attachment. Dried peridium brownish orange (5C-4-6) to light brown (6D-4, 5),
mottled with dark brown (6F-8); rhizomorphs darker than peridium, but not
245
conspicuous. GLEBA: Loculate, locules as dried small, 5-8 per mm, only partially
filled at maturity; white when young, then buffy. Dried gleba brownish orange (5C-4-6)
ODOR fragrant in older sporocarps. KOH on dried peridial surface quickly dark
brown; merely darker on the gleba. FeSOq on dried peridial surface negative; merely
darker on the gleba.
SPORES: 7.5-9.0 x 3.0-4.0 pm, oblong to short cylindrical, often irregularly
bent, truncate, basal scar and sterigmal remains prominent, walls thickened at maturity,
smooth, in KOH, pale yellowish brown singly and in mass; pale yellow in Melzer's
reagent, golden yellow in mass, mostly with 2 lipid droplets per spore. HYMENIAL
ELEMENTS: Basidia borne in hymenial layer, clavate, 10-20 x 5.5-8 pm, collapsed as
dried, mostly 4- or 8-spored; brachybasidioles 20-30 x 10-15 pm, thin-walled in youth,
thick-walled, mucilaginous in age, not easily disarticulated in crush mounts, heavily
gelatinized. GLEBA: Trama of hyaline, cylindrical, hyphae that are 3-5 pm diam,
loosely interwoven near the hymenium, subparallel in the mediostratum, mucilaginous
with age, highly refractive in KOH. Subhymenium cellular, not well defined, 1-2 cells
deep, cells 5-10 pm diam, globose to subglobose, hyaline, thick-walled with age.
PERIDIUM: 300-450 pm; of loosely interwoven, cylindrical but knobby, swollen,
highly branched, moderately thick-walled hyphae, 3-5 pm diam; hyphal strands common,
scattered throughout, 10-25 pm diam, 3-8 cells thick, twisted into small knots of hyphae;
oleiferous hyphae common, scattered throughout, 3-5 pm diam, yellowish refractive in
KOH and Melzer's reagent, cylindrical, knobby or contorted and irregularly swollen; in
KOH, pale yellow, the pigment diffusing into the fluid; encrusted with yellowish
refractive pigment masses in Melzer's reagent. Subcutis thin or absent, composed: of
highly gelatinous or oleiferous hyphae. Clamp connections absent.
HABITAT AND SEASON: Hypogeous just below the surface of the humus
in low damp pine woods. December.
COLLECTIONS EXAMINED: HOLOTYPE: South Carolina, Hartsville, Coker 5999
(NCU).
I have not collected this fungus fresh. The spores resemble R. truncatus in shape
and overall morphology but are narrower and not dextrinoid at any point in development.
The length of the sterigmal remains on the spores is also consistently longer. The
peridium of R. atlanticus is similar to that of R. truncatus but is much thicker.
Doubtful Species of Rhizopogon in the southeastern United States
R. parasiticus Coker and Totten, Jour. Elisha Mitchell Sci. Soc. 39:101-109.
The specimens at NCU (3797, 3990, 5383, 6051, 7589) and at NY (6057) are no longer
in any state to be examined; they are composed solely of soil and root fragments. From
the original description, however, these are nothing more than large tuberculate
ectomycorrhizae common throughout the southeastern United States. I have found
senescent forms of these ectomycorrhizae late in the season that are internally parasitized
by different corticioid basidiomycetes. I believe that the spores found by Coker and
Totten are those of a parasite, not of a parasitizing Rhizopogon species.
R. roseolus (Corda) Hollos, in Coker and Couch, Gasteromycetes of Eastern
United States and Canada, p. 32-33. Only two collections originally cited by Coker and
Couch under this name could be located at NCU, 7212 and 7214. Collections 7207 and
7229 could not be found. Both collections 7212 and 7214 are young specimens of R.
rubescens (Tul.) Tulasne.
R. maculatus Zeller and Dodge, in Coker and Couch, Gasteromycetes of
eastern United States and Canada, p. 35. Collection 955 (NCU), which was included
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218
by Coker and Couch under this name, could not be located. I am inclined to believe that
this species does not occur in our area and that another fungus, possibly R. vulgaris, is
the one described by Coker and Couch.
R. hesleri Dodge, "Checklist of Fungi of the Great Smoky Mtns. Nat. Park,"
Management Report no. 29, p. 95. I find no validly published Rhizopogon by this
name. Collection 6609 (TENN), rightly identified by Dr. Smith as R. vulgaris, contains
a note from Dodge indicating that he was designating it as the type of R. hesleri.
Likewise, collection 4860 (TENN), a collection of R. couchii, contains an annotation
from Dodge identifying it as R. hesleri.
ACKNOWLEDGMENTS
The work presented in this study would not have been possible without the tremendous
contributions of Dr. Alexander Smith, to whom I am dedicating this paper. Thanks also
to Drs. James Trappe and David Hosford for their constructive comments on this
manuscript.
LITERATURE CITED
Coker, W. C., and J. N. Couch. 1928. The Gasteromycetes of the Eastern United
States and Canada. Univ. N. Carolina Press, Chapel Hill, N. Carolina.201 p.
Hosford, D. R. 1975. Taxonomic studies on the genus Rhizopogon I. Two new
species from the Pacific northwest. Nova Hedwigia 51: 163-169.
, and J. M. Trappe. 1980. Taxonomic studies in the genus
Rhizopogon. Il. Notes and new records of species from Mexico
and Caribbean countries. Bol. Soc. Mex. Mic. 14: 3-15.
Kornerup, A., and J. H. Wanscher. 1981. Methuen Handbook of Colour. 3rd ed. Eyre
Methuen, London, 252 p.
Lanjouw, J., and F. A. Stafleu. 1964. Index Herbariorum part I. The Herbaria of the
World. Regnum Vegetabile 31: 1-251.
Linder, D. H. 1924. A new species of Rhizopogon from New Hampshire. Rhodora.
26: 196-197.
Smith, A. H., and S. M. Zeller. 1966. A preliminary account of the North American
species of Rhizopogon. Mem. N.Y. Bot. Gard. 14: 1-178.
Svrcek, M. 1958. Rhizopogon. In Flora CSR I. Gasteromycetes, A. Pilat et al., eds,
Praha.
Trappe, J. M. 1975. A revision of the genus Alpova with notes on Rhizopogon and
the Melanogastraceae. Nova Hedwigia 51: 279-309.
Tulasne, L. R., and C. Tulasne. 1844. Fungi nonnulli Hypogaei, Nowi vel minus
cogniti. Parlatore, Giorn. Bot. Ital. 2: 58.
Zeller, S. M., and C. W. Dodge. 1918. Rhizopogon in North America. Ann. Mo.
Bot. Gard. 5: 1-36.
MYCOTAXON
VOLPOXXV ID epp 219-255 October-December 1986
SPECIES OF SCUTELLOSPORA (ENDOGONACEAE) WITH SMOOTH-
WALLED SPORES FROM MARITIME SAND DUNES: TWO NEW SPECIES
AND A REDESCRIPTION OF THE SPORES OF SCUTELLOSPORA
PELLUCIDA AND SCUTELLOSPORA CALOSPORA
R. E. KOSKE
Department of Botany, University of Rhode Island, Kingston,
Rhode Island 02881, USA
and
CHRISTOPHER WALKER
Forestry Commission, Northern Research Station, Roslin, Midlothian, EH25 9SY,
UK
SUMMARY
Examination of soil samples from sand dunes on the eastern seaboard of the USA revealed two
undescribed species of Scutellospora with smooth outer walls. Scutellospora weresubiae is
characterized by its translucent pink spores with a complex wall structure, and S. fulgida
produces hyaline spores with three walls. The species are described in this paper, and
comparisons drawn with two other similar smooth-walled species, S. pellucida and
S. calospora, which are illustrated and redescribed with a standardized wall terminology.
A survey of the vesicular-arbuscular mycorrhizal (VAM) fungi associated with the
principal dune-colonizing plants of the east coast of the United States revealed two
undescribed species of Scutellospora Walker & Sanders with smooth outer walls. The
recent recognition of the taxonomic significance of the details of spore wall structure
in the Endogonaceae prompted us to re-examine other species of Scutellospora with
smooth-walled spores for comparison with the new species. We redescribe the spores
of two previously described species, applying a standardized wall nomenclature.
MATERIALS AND METHODS
Spores were recovered from dune samples by a wet-sieving and filtration technique
(Koske and Walker 1984). Specimens were mounted on microscope slides in a
polyvinyl alcohol mountant with (PVL) or without (PVLG) lactophenol (Koske &
Tessier 1983; Walker 1979), or, in the case of those type specimens preserved in
lactophenol (S. calospora (Nicol. & Gerd.) Walker & Sanders and S. pellucida
(Nicol. & Schenck) Walker & Sanders), in PVL, PVLG or lactophenol. Because
parasitism by soil micro-organisms destroys inner wall structures, only apparently
healthy spores were described. Collection numbers in descriptions, preceded by
‘Koske’ refer to soil sample collections and those preceded by ‘Walker’ to herbarium
accession numbers. Type material was kindly loaned to us by the curator of the
Farlow Herbarium.
220
221
Wall descriptions and terminology are according to Walker (1983, 1985) and Morton
(1986a). Although the details of wall structure described in this study can be observed
by brightfield illumination with a good-quality microscope properly illuminated and
adjusted, the smaller features are more easily observed with differential-interference
contrast (Nomarski) techniques. Colors are designated where possible by the color
chart published by the Royal Botanic Garden, Edinburgh (Anon 1969).
Holotypes have been deposited in the herbarium at Oregon State University (OSC).
Isotypes have been placed in the Farlow Herbarium (FH), and at Kew (K).
RESULTS AND DISCUSSION
SCUTELLOSPORA FULGIDA Koske et Walker sp. nov. Figs. 1A-G & 5
Sporae in solo singillatim efformatae, globosae vel subglobosae, 160-240 x 160-245 um, lucentes, hyalinae
vel straminellae. Tunicae sporae stratis tribus in termis duabus. Turma externa stratis duobus: stratum
extimum laeve, fragile, hyalinum, usque ad | 4m crassum; stratum internum lamellatum, hyalinum, ad
stratum externum arcte haerens, 6-8 um crassum, in solutione Melzeri pallide aurantiascens. Turma
interna strato uno, hyalino, membranaceo, 0.5-1 um crasso. Cellula suspensoriformis hepatica, distincte
fuscior quam spora, 26-46 wm lata, tunica 1-2(-2.5) wm crassa. Cellulae auxiliares nodosae 6-10 in
fasciculo, hyalinae vel pallide hepaticae, irregulares, 20-25 x 25-37 um, prominentiis plerumque truncatis,
conicis, vel subcylindricis, nodosis, 2-10 x 2-10 um.
Spores formed singly in the soil, terminally on a bulbous suspensor-like cell;
glistening, hyaline to pale straw or pale yellowish-cream; globose to subglobose;
160-240 x 160-245 um.
Spore wall structure (see murograph, Fig. 5) of three walls (walls 1-3) in two groups
(groups A and B). Group A with an outer smooth, brittle, hyaline unit wall (wall 1) up
to | um thick, tightly adherent to an inner, brittle, hyaline, laminated wall (wall 2)
6-8 um thick. Group B of a single, thin (0.5-1 wm), membranous wall enclosing the
spore contents. Wall 2 turning pale orange in Melzer’s reagent, other walls not
reacting.
Germination shield oval, 40-90 x 80-130 4m formed by the membranous wall.
Suspensor-like cells borne terminally on a sparsely septate subtending hypha;
26-46 um broad, yellow-brown, distinctly darker in color than the spore, with walls
1-2(-2.5) um thick, bearing one to several stout projections 6-38 x 2-12 um.
Fig. 1: Scutellospora fulgida.
Whole spore, x 200.
Suspensor-like cell with hyphal projections, x 500.
Crushed spore. Wall 3 (arrow) has separated from wall group A, x 160.
Germination shield. Margin is indicated by arrows, x 485.
Wall structure of a crushed spore. Wall one has fractured and folded back
from wall 2. Wall 3 is indicated, x 1235.
F. Auxiliary cells, x 620.
G. Cluster of auxiliary cells, x 520.
MOOD > ;
2o2
Auxiliary cells in soil borne on coiled, hyaline hyphae 4-5 um diameter in clusters of
6-10(-20), hyaline to pale yellow-brown, irregularly shaped, 20-25 x 25-37 um, with
broad, usually truncated, conical to subcylindric knobby projection 2-10 x 2-10 um.
DISTRIBUTION AND HABITAT: Known only from a limited area of coastal sand
dunes on the eastern North American seaboard. Scutellospora fulgida occurred
frequently in samples taken from maritime dunes at the southern tip of New Jersey
and farther south to Maryland and Virginia. However, it was not recovered from any
of more than 500 samples collected from mid New Jersey to as far north as Maine.
MYCORRHIZAL ASSOCIATIONS: Found in the root zones of Ammophila
breviligulata Fern., Solidago sempervirens L., and Uniola paniculata L., but not
proven mycorrhizal.
ETYMOLOGY: Latin, fulgida, (shining, gleaming, glittering), referring to the
appearance of the spores when viewed through the dissecting microscope.
COLLECTIONS EXAMINED: HOLOTYPE: VIRGINIA - Virginia Beach Township, Seashore State
Park, among roots of A. breviligulata on sand dunes (Koske 505; Walker 658; 23 April 1983). Other
Collections, NEW JERSEY - Cape May Co., Cape May Point Sand Dunes (Koske 401, 404; 14 March
1982); VIRGINIA - Accomack Co., Assateague Island Sand Dunes (Koske 437, 440, 445, 457; 15 March
1982).
Wall | of Scutellospora fulgida is usually easy to observe in crushed spores (Fig. 1 F),
but in some specimens it is thin and difficult to see, even under high magnification.
Wall 3 also can be difficult to detect as it may be tightly adherent to wall 2.
Scutellospora fulgida differs from other species mainly by its hyaline spores and
relatively simple wall structure. It most resembles S. pellucida or hyaline specimens
of S. calospora when viewed with a dissecting microscope. The inner wall group
(group B) of S. fulgida spores is thin, consisting of a single membranous wall
(Figs. 1C, F). The formation of the germination shield in spores of S. fulgida may lead
to the erroneous impression that group B has two walls. The inner wall group of
S. calospora consists of two juxtaposed membranous walls (Figs. 4D, F), while that
of S. pellucida consists of three walls (Fig. 3G). In addition, the innermost wall of
both S. calospora and S. pellucida spores becomes red or reddish-purple in Melzer’s
reagent, while that of S. fulgida stains yellow. Spores of all three of these species
possess a thin outermost unit wall. Spores of S. fulgida develop a pale yellow
coloration in the walls upon storage in polyvinyl alcohol mountants and thus may be
confused with colored spores of S. calospora, but the different wall structures and the
formation of clustered auxiliary cells in the former (Figs. 1E, G) compared to single
auxiliary cells in the latter (Figs. 4H, I) serve to separate the species under such
circumstances.
Scutellospora fulgida could, on examination under a dissecting microscope, also be
mistaken for S. gilmorei (Trappe & Gerd.) Walker and Sanders, another hyaline-
spored species. Examination of mounted specimens through a compound
microscope reveals, however, that although spores of the latter possess an outer wall
group similar to S. fulgida, their inner wall group is composed of four easily separable
walls rather than a single membranous wall.
223
224
Another hyaline-spored species, S. savannicola (Ferr. & Herr.) Walker & Sanders,
has been described from Cuba. We have been unable to obtain specimens of the type
material to examine, but the published description (Ferrer and Herrera 1981) is of a
spore with five walls in three groups, an outer wall group composed of two walls up to
3 wm and 2.3 um thick, respectively, a middle wall group of two membranous walls,
1.5 and 1.3 um thick, respectively, and an inner wall | um thick.
SCUTELLOSPORA WERESUBIAE Koske et Walker sp. nov. Figs. 2A-F & 5
Sporae in solo singillatim efformatae, globosae, subglobosae vel irregulares, 125-265 x 135-414 um,
lucentes, pallide vel saturate incarnatae. Tunicae sporae stratis sex in turmis trilus. Turma externa stratis
duobus: stratum extimum laeve, fragile, hyalinum, usque ad 0.5 um crassum, adstratum internum, fragile,
incarnatum, 3-8.8 um crassum, in solutione Melzeri interdum rubescens haerens. Turma media stratis
duobus membranaceis, utrumque usque ad | um crasso. Turma interna strato extimo coriaceo, hyalino,
2-8 um crasso, stratum membranaceum, hyalinum, 0.5 um crassum, in solutione Melzeri rubescens,
cingenti. Cellula suspensoriformis hyalina vel pallide testacea, 32-50 um lata, tunica distaliter | wm crassa,
prope sporam usque ad 5 wm. Cellulae auxiliares nodosae 5-14 in fasciculo, incarnatae vel pallide brunneo,
subglobosae, 17-33 x 25-40 um, prominentiis plerumque truncatis, conicis nodosis, 4-7 x 7-14 wm.
Spores formed singly in the soil terminally on a bulbous suspensor-like cell;
translucent, glistening, pale pink to deep pink; globose to subglobose or irregular;
125-265 x 135-414 (mean 210 x 210) um.
Spore wall structure (see murograph, Fig. 5) of six walls (walls 1-6) in three groups
(groups A, Band C). Group A often with an outer, smooth, brittle, hyaline, unit wall
(wall 1) up to 0.5 wm thick tightly adherent to an inner brittle, pink, laminated wall
(wall 2) 3-8.8(-15) um thick that sometimes turns red in Melzer’s reagent.
Group B of two membranous walls (walls 3 and 4), each up to | wm thick, wrinkled in
crushed spores.
Group C formed by a thick hyaline coriaceous wall (wall 5) (2-8 um thick)
surrounding a hyaline membranous innermost wall (wall 6) 0.5 um thick that turns
red in Melzer’s reagent.
Germination shield, known only from two specimens, cardioid, 100 x 69-80 um with
a smooth margin | wm thick and with a germ tube initial in each lobe.
Suspensor-like cells borne terminally on a sparsely septate or aseptate subtending
hypha, 32-50 wm broad, hyaline to pale brownish-yellow, with walls 1 um thick
Fig. 2. Scutellospora weresubiae.
A. Whole spore, x 140.
B. Germination shield with germ tube initial (arrow), x 365.
C. Crushed spore in Melzer’s reagent. Wall 6 is darkly stained. Wall 5 is not
visible in this photograph. Walls 3 and 4 are wrinkled, x 125.
D. Wall structure. Walls one and 6 are not easily visible in this photograph.
Walls 3 and 4 are closely associated. Note wrinkling of wall 3 (arrow) x 365.
E. Composite photograph of suspensor-like cell with hypal pegs, x 495.
F. Auxiliary cells, x 480.
ZLo
distally, thickening to 5 um at the spore base. One or two hyphal pegs up to 27 um
long x 3-8 wm wide may arise from the suspensor-like cell and project towards the
spore base. Pegs not occurring on all specimens.
Auxiliary cells in soil borne in clusters of 5-14, bright pink when fresh, fading to pale
brown; subglobose, 17-33 x 25-40 um, with blunt, rounded knobby projections 3-6 x
4-7 um or broad truncated conical projections 4-7 x 7-14 um; produced on coiled
pink hyphae 3-4 wm diameter.
DISTRIBUTION AND HABITAT: Known only from coastal sand dunes in
Virginia, Florida and South Carolina, and from sandy spoil heaps of acopper mine in
Malaysia.
At first we thought S. weresubiae to be limited in its distribution to an area stretching
from the south end of the Assateague Island, southward to South Carolina.
Searching for the species among 21 samples from the north end of the 60 km-long
island, from 41 samples among dunes in New Jersey and from more than 400 samples
from dunes of the Rhode Island and Massachusetts coast, proved fruitless, despite
the fact that the vegetation and edaphic conditions in all these areas are similar.
However, the species later was identified from collections in Florida (D. Sylvia, pers.
comm.) and from Malaysia (G. Sankaralingam, pers. comm.). Both these last
collections were from sand, and it seems that the species is climatically and
edaphically controlled and might be expected to occur in sandy conditions in
relatively warm areas.
MYCORRHIZAL ASSOCIATIONS: Scutellospora weresubiae was found in
association with field-grown plants of Ammophila breviligulata, Uniola paniculata,
and Solidago sempervirens, but failed to form mycorrhizae in pot culture with onion
(Allium cepa L. cv Agway Sweet Spanish), clover (Trifolium repens L.), and corn
(Zea mays L. cv Seneca).
ETYMOLOGY: Named in honor of Luella K. Weresub, distinguished mycologist.
COLLECTIONS EXAMINED: HOLOTYPE: VIRGINIA - Accomack Co., Assateague Island sand
dunes from underneath Solidago sempervirens (Koske 440, Walker 636; 15 March 1982). Other
collections: VIRGINIA -Accomack Co., Assateague Island (Koske 438, 439, 445, 459; 15 March 1982),
(Koske 294; 20 April 1981); Virginia Beach Township, Seashore State Park (Koske 502, 506, 522, 526; 23
March 1983); SOUTH CAROLINA - Horry Co., Myrtle Beach (Koske 288; 14 April 1981).
The complex wall structure and pink color of S. weresubiae spores distinguish them
from other species. The outermost wall (wall 1) may be difficult to observe, and can
best be seen on specimens that have been vigorously crushed. The pink wall (wall 2)
loses its color almost immediately when spores are placed in lactophenol, formalin,
or polyvinyl alcohol, although it is retained for a few days in glycerol jelly mounts. In
fresh specimens, wall 6 quickly stains dark red in Melzer’s reagent. Wall 2 usually
stains red also, but only after a longer time. Occasionally spores possess one or two
laminae broken loose from wall 2. These may be misleading, appearing as additional
unit walls between walls 2 and 3. The walls of the inner wall group may best be
observed by allowing crushed spores to stain in Melzer’s reagent for about 15 minutes
and then adding a polyvinyl alcohol mounting solution to the edge of the coverslip
226
and allowing it to infiltrate overnight or by crushing spores in PVL or PVLG to which
has been added a drop of freshly prepared Melzer’s reagent. The inner wall group is
difficult to interpret. In wall group B, walls 3 and 4 loosely surround walls 5 and 6 of
group C and frequently are noticeably wrinkled (Figs. 2C, D). Wall 3 is thin
(<0.2 um) and may seem absent from some spores. It is, however, probably present in
all spores, but may be so thin that it cannot be resolved by light microscopy. Studies
of cracked spores by scanning electron microscopy confirmed the presence of this
wall on all specimens examined, along with the other walls observed by light
microscopy. Spores fixed in formaldehyde often show wall 3 and 4 associated with
wall group A. In unfixed specimens they are usually close to walls 5 and 6 in Group C.
We therefore placed them in a group of their own to avoid confusion. Wall 5 stains
yellow in Melzer’s reagent and is pliable, although up to 8 wm thick. Wall 6 is best
seen by its strong reaction to Melzer’s reagent, this wall and wall 3 often being
extremely difficult to observe if unstained.
The color of S. weresubiae spores fades with age, and many older specimens retain
only a faint pink tinge. The bright pink color of fresh spores was not reduced by
immersion for several days in phosphate buffer at pH 4 or pH 10. In water, the color
pales considerably after a day or two, though it usually still can be detected when the
spores are observed with a mixture of transmitted and reflected light. Spores of
S. weresubiae, incubated on filter paper placed on moist sand, germinated after two
weeks to produce bright pink germ tubes, hyphae, and auxiliary cells. In preservative,
such as formaldehyde, gluteraldehyde, formalin-acetic acid, or lactophenol, the color
is lost very rapidly. Thus, it is important that the color of spores is recorded when they
are freshly extracted from the soil.
S. weresubiae resembles S. alborosea (Ferr. & Herr.) Walker & Sanders, a pink-
spored species reported only from Cuba, although the wall structure of the two
species seemingly differs substantially. Whereas the former has spores with six walls
in three groups, the latter is described as having only two wall groups, the outer group
with only two walls and the inner with a single membranous wall. We have
unfortunately been unsuccessful in obtaining material of S. alborosea for
examination and have thus had to make this comparison from the description in the
protologue.
The only other pink-spored species in the Endogonaceae is Gigaspora rosea Nicol. &
Schenck. Spores of this species are opaque and creamy-white with a coral-pink
coloration near the suspensor-like cell. They are not translucent and do not have the
overall bright pink color of spores of S. weresubiae. In addition, spores of G. rosea
have the simple wall structure, direct germination characteristics and echinulate
auxiliary cells that distinguish Gigaspora spp. from species of Scutellospora (Walker
& Sanders 1986).
Very lightly pigmented or faded spores of S. weresubiae resemble those of
S. pellucida, and spores of both species have six walls in three groups. Wall | in both
is a unit wall, closely appressed to the laminated second wall, but wall | of the former
is thin (up to 0.5 wm) and often extremely difficult to see, whereas the corresponding
wall of the latter is relatively thick (1-2 4m) in most specimens and can normally be seen
228
clearly at 400 x magnification (Figs. 3C, E). The structure of the inner wall groups
(Group B and C) of the two species may appear similar at low magnification.
However, wall group B of S. pellucida consists of a very thin membranous wall and
two unit walls, with the membranous wall closely adherent to one of the unit walls
(Figs. 3F, G). In S. weresubiae, wall group B is comprised of two membranous walls
(Figs. 2C, D), each usually thicker than the membranous wall of S. pellucida.
Group C of S. pellucida is an amorphous wall (Figs. 3F, G) that has plastic properties
in PVL and PVLG (see S. pellucida discussion) and stains reddish-purple in Melzer’s
reagent. Wall group C of .S. weresubiae is formed by a thick, coriaceous wall and a
thin, membranous innermost wall (Figs. 2C, D) that stains red (but not reddish-
purple) in Melzer’s reagent. The taxonomic significance of such staining differences
in some Endogonacea has been discussed by Morton (1986b).
The differences in wall structure of S. weresubiae and S. pellucida can be compared in
the murographs (Fig. 5).
REDESCRIPTION OF SCUTELLOSPORA PELLUCIDA AND
S. CALOSPORA
In comparing the newly described species of Scutellospora with other smooth-walled
species, it became clear that a redescription of spores of certain existing taxa making
use of the standardized wall terminology suggested by Walker (1983), may be helpful.
The following descriptions of the spores of S. pellucida. and S. calospora are based
upon examination both of type specimens and other collections. Specimens of
S. pellucida from sand dunes in Virginia (Koske 518) were up to 250 x 410 um;
somewhat larger than originally described, but we consider this to be intraspecific
variation. The size range of S. calospora has also been increased from measurements
of several collections.
SCUTELLOSPORA PELLUCIDA (Nicol. & Schenck) Walker & Sanders
Figs. 3A-H & 5
Spores formed singly in the soil or in roots, borne terminally on a bulbous suspensor-
like cell; glistening with oil droplets, hyaline to pale grey, globose, ellipsoid, or
irregular; 58-183(-250) x 58-241(-410) um.
Fig. 3: Scutellospora pellucida.
A. Whole spore, x 235.
B. Suspensor-like cell, x 570.
C. Crushed spore showing wall groups A and B. Wall one (arrow) has
separated from wall 2, x 140.
D. Germination shield. Margin (arrow) with invaginations, x 235.
E. Wall structure of spore preserved in formalin. Note walls 1 and 2, and fused
walls of group B, x 935. ;
F. Wall structure of fresh spore in PVLG. Wall one is just visible (arrow). Note
separation of wall 6 from other walls, x 1310.
G. Wall structure of fresh spore crushed in Melzer’s reagent. Walls 3 and 4,
typically, are closely adherent in this spore. Wrinkling of wall 3 is indicated (W).
Note dark staining of wall 6, x 870.
H. Auxiliary cells, x 350.
Zag
Spore wall structure (see murograph, Fig. 5) of six walls (walls 1-6) in three groups
(groups A, B and C). Group A with an outer smooth, brittle, hyaline, unit wall (wall
1) 1-2 um thick, closely appressed to an inner brittle, hyaline, laminated wall (wall 2)
2-7 (-18) wm thick. Group B consisting of a hyaline membranous wall (wall 3) (1 wm
thick) closely adherent to two hyaline unit walls (walls 4 and 5). Wall 4, 1-2 um thick;
wall 5, 0.8-3 wm thick. Group C consisting of a single, hyaline amorphous wall (wall
6), 2-5 wm thick when measured in Melzer’s reagent, of variable thickness in PVL or
PVLG (see below). Walls 2 and 5 typically stain reddish-purple in Melzer’s reagent,
although wall 2 remains unstained in some specimens.
Germination shield oval, 110-140 x 125-160 um, the margin with invaginations.
Suspensor-like cell borne terminally on a septate subtending hypha; 10-29(-50) um
broad, hyaline; walls <0.5 um thick distally, thickening to 2.5 um near the spore base.
COLLECTIONS EXAMINED: USA: FLORIDA - Jay (Isotype in FH); CALIFORNIA - Santa Barbara
Co., San Miguel Island Dunes (Koske 561, 567-570, 572, 575, 576, 588; 5 July 1984); MASSACHUSETTS
- Bristol Co., S. Dartmouth Dunes (Gemma 2-20-6; February 1985); NEW JERSEY - Cape May Co., Cape
May Pt. Dunes (Koske 401, 407, Walker 519; 14 March 1982); Ocean Co., Ocean City Dunes (Koske 382;
13 March 1982); RHODE ISLAND - Washington Co., Moonstone Beach Dunes (Friese P3, Koske A7;
6 August 1983); SOUTH CAROLINA - Horry Co., Myrtle Beach Dunes (Koske 288; 14 April 1981);
VIRGINIA - Virginia Beach Township, Seashore State Park Dunes (Koske 505, 518, 520, 23 April 1983);
WEST VIRGINIA - Preston Co., pot culture from mine site (Morton 396, Koske A8).
The outermost unit wall (wall 1) of S. pellucida was easily resolved in most specimens,
sometimes separating distinctly from wall 2 (Figs. 3C, E), though in some specimens
it was not clearly distinguishable (Figs. 3G, H). The laminated wall (wall 2) of spores
of S. pellucida typically is 2-7 um thick, but in a few spores from high-aluminum,
low-pH sites in West Virginia, this wall reached a thickness of up to 18 um.
Dimensions of other walls in these samples matched those of more typical specimens.
The inner wall groups (B and C) of S. pellucida can be very difficult to interpret,
especially at lower magnifications when they may appear as a single, hyaline, flexible
wall 1.5-5 um thick with 2-4 apparent laminations. The appearance and thickness of
the walls in groups B and C depends upon the state of the spores when the slide was
made (fresh or preserved), the mounting medium, and how the spores were crushed
(once or twice). When fresh spores are crushed in Melzer’s reagent, wall 6 stains dark
reddish-purple within a minute, and walls 3-5, which remain unstained, can often be
distinguished (Fig. 3G). Frequently, however, wall 3 is so closely appressed to wall 4
that often it is detectable only by the fine wrinkling it exhibits (Fig. 3G). In some
specimens, walls 3-5 remain so close together that they appear as a single wall.
When spores preserved in 5% formaldehyde for more than a few months were
crushed in Melzer’s reagent, the distinctive staining reactions of walls 2 and 6 did not
occur. Wall 2 became reddish-brown, and wall 6 did not stain. A similar lack of
staining in Melzer’s reagent following preservation occurs in spore walls of several
species of Acaulospora (Morton 1986b). A second change that occurred when spores
of S. pellucia were preserved was the fusion of walls 3-6 to form a single, thick,
apparently laminated wall (Fig. 3E).
230
_
Zon
Fresh spores of S. pellucida mounted in PVL or PVLG and crushed once exhibit a
wall structure generally similar to that of spores mounted in Melzer’s reagent (Fig.
3F). Wall 6 may appear slightly wrinkled in some spores that have been so treated. If,
after a few minutes, pressure is again applied to the coverslip, the amorphous nature
of wall 6 becomes apparent, and the wall becomes coarsely wrinkled. The interval
between crushings apparently allows the PVL or PVLG to interact with the wall,
giving it a plastic consistency (Morton 1986a, b). If sufficient force is applied, wall 6
may deform markedly and extrude completely from wall group B. The amorphous
innermost wall of spores of some Acaulospora species may be similarly extruded
when spores are crushed (Morton 1986a). The thickness of wall 6 when measured
after the second crushing thus is dependent upon the pressure exerted during that
crushing. For this reason, the thickness range given for this wall (2-5 um) is based on
specimens mounted in Melzer’s reagent which does not induce the plasticity. The
amorphous wall type, its staining reactions and other characteristics were originally
defined and described by Morton (1986a, b) for a species of Acaulospora.
The variable appearance of wall 6 as influenced by slide preparation techniques and
freshness of the spores, and the difficulty in resolving the 3 walls of group B make
S. pellucida one of the most difficult species to identify.
SCUTELLOSPORA CALOSPORA (Nicol. & Gerd.) Walker & Sanders
Figs. 4A-I & 5
Spores formed singly in the soil, terminally on a bulbous suspensor-like cell;
translucent, hyaline to pale greenish-yellow; globose, ellipsoidal or cylindrical,
occasionally broader than long; 114-285(-511) x 110-412(-511) um.
Spore wall structure (see murograph, Fig. 5) of four walls (walls 1-4) in two groups
(groups A and B). Group A consisting of an inner, brittle, hyaline to pale yellow, very
finely laminated wall (wall 2) 3-5 um thick that may be surrounded by a thin, very
closely appressed hyaline unit wall (wall 1), 0.5-1 wm thick. Group B of two hyaline
membranous walls (walls 3 and 4). Wall 3, 0.5-1 um thick, often wrinkling in crushed
spores. Wall 4, 1-1.5 «wm thick, staining red in Melzer’s reagent.
Fig. 4: Scutellospora calospora.
A. Whole spore, x 200.
B. Suspensor-like cell, x 470.
C. Crushed spore showing wall groups A and B, x 115.
D. Crushed spore. Wall 3 has separated from wall 4. Wall one is not visible,
x 470.
E. Germination shield in face view. Margin indicated by arrows, x 345.
F. Spore wall structure. Walls 2-4 are indicated. Note laminations in wall 2.
Wall one is not apparent, x 1150.
G. Germination shield, lateral view. Note location of suspensor-like cell
(arrow), X 395.
H. Auxiliary cell with smooth surface, x 415.
|. Auxiliary cells with Knobby surface, x 415.
Zz
SE NINN
-Raansys 32
Gigaspora calospora
Gigaspora fulgida
Gigaspora pellucida
Gigaspora weresubiae
Big e)
Germination shield oval, 35-70 x 50-90 um, often with invaginations along the
margin.
Suspensor-like cell borne terminally on a septate subtending hypha; 33-48 um broad;
walls concolorous with wall 2 of spore; walls 1 um thick distally, thickening
somewhat near the spore base.
COLLECTIONS EXAMINED: UK: SCOTLAND - Perthshire, East Newton (Holotype in FH);
SCOTLAND - Midlothian, Northern Research Station of the Forestry Commission, in pot culture with
Trifolium repens L. started with a single spore from a grassy verge in the grounds of the research station
(Walker 1002; 13 February 1985). ENGLAND -Hertfordshire, Rothamsted Experiment Station, from a
pot culture (with Allium porrum L.) the origins of which are unknown (Walker 1153; 22 August 1985).
USA: CALIFORNIA - Santa Barbara Co., San Miguel Island Dunes (Koske 568, 581, 596; 5 July 1984);
MASSACHUSETTS - Suffolk Co., Plum Island Dunes (Koske 339; 9 October 1981); NEW JERSEY
-Monmouth Co., Spring Lake Dunes (Koske 361; 13 March 1982); RHODE ISLAND - Washington Co.,
Moonstone Beach Dunes (Koske 278; 18 February 1981: Friese 63; 8 April 1983).
The characterization of Scutellospora calospora is made somewhat difficult by the
paucity and poor state of preservation of the type material, which consists of a few
spores mounted (by Walker in 1979) in PVL and three or four spores originally
preserved in lactophenol. Hall and Abbott (1984) examined this material and
redescribed the species. They indicated a wall structure which, in the terminology of
Walker (1983), has an outer wall group (A) consisting of a thin unit wall surrounding
a laminated wall, and an inner wall group (B) of a single membranous wall. We have
examined the type material and have drawn different conclusions. The outer wall
often is very difficult to see in most specimens, though we agree it probably is present,
and we are in complete accordance with Hall and Abbott about wall 2, a laminated
wall to which wall | is tightly adherent. We differ in our interpretation of the inner
wall group. In all the specimens we examined, we observed that wall group B
consisted of two adherent membranous walls (Figs. 4D, F).
There is some confusion in the literature regarding the color of spores of S. calospora.
In recent years, the species has been considered to have yellow or yellow-green spores,
and the impression has been given that only immature spores are hyaline. Gerdemann
and Trappe (1974) described the species as having ‘pale yellow to greenish yellow’
spores, and Trappe (1982) in his synoptic key included S. calospora only in the color
group with ‘outer wall layer yellow’. Hall and Abbott (1984) did not describe the color
of the spores, but they did note that the ‘middle wall layer’ was ‘coloured’. Ina key to
the species of Endogonaceae (Hall and Fish 1979), S. calospora was described as
having ‘mature spores pale yellow to pale greenish yellow’, and in a later key the
species was similarly included only in couplets derived from spores that are ‘coloured
Fig. 5: Murographs of four species of Scutellospora. Shading is by vertical
alternating dotted lines for laminated walls, 45° hatching in one direction for
membranous walls, 45° hatching in both directions for coriaceous walls, and
curved lines for ‘amorphous’ walls. Unit walls are left unshaded. Walls marked
with an asterisk are very difficult to see.
234
though may be hyaline when young’ (Hall 1984). The protologue of Scutellospora
calospora describes the spores as ‘colorless or light yellow’ (Nicolson and Gerdemann
1968), and in a key to the Endogonaceae of Florida, Nicolson and Schenck (1979)
indicate that they are ‘hyaline, white or shades of yellow’.
This confusion was compounded when we isolated (in both single-spore and multi-
spore pot cultures) a small spored Scutellospora from Scotland that had hyaline or
white spores of the correct size range of S. calospora and possessed a wall structure
exactly coinciding with our interpretation of the type material of that species (though
sometimes wall 4 was extremely thin and could be detected only when fresh spores
were treated with Melzer’s reagent). Under storage in moist soil, or when preserved in
5% formaldehyde solution, spores from our cultures generally remained hyaline,
although after a few days in water at room temperature, some of them became a little
yellowish (though it should be emphasized that this coloration was extremely pale,
and easily noticeable only in comparison with freshly extracted specimens). When
stored in lactophenol, however, the spores quickly became yellow to yellow-olive.
To resolve these questions of spore color, we sought the advice of Dr Tom Nicolson,
one of the original authorities for Scutellospora calospora, who kindly agreed to
examine our culture material. He concluded that our collections agreed with his
concept and memory of the species, and stressed that the material he described was
mainly colorless, but that some specimens appeared slightly greenish-yellow. In view
of this, we consider that S. calospora has hyaline or white to very pale greenish-yellow
spores. It seems likely that dark yellow or greenish-yellow spored species that may
have been attributed to S. calospora might more properly be placed as S. aurigloba
(Hall) Walker and Sanders or some other taxon. Nevertheless, we are unsure as to
how much significance to give to spore color as a taxonomic criterion, and if yellow
spores are found that have the size range, wall structure, germination characteristics,
and auxiliary cell features typical of S. calospora, then consideration should be given
to extending the species concept to include them in that taxon, rather than to
describing a new species. In view of this uncertainty, any use of isolates considered to
be this fungus should be backed up by a careful description of the spores, deposition
of specimens in a herbarium, and where possible, preservation of living cultures for
future examination and use.
ACKNOWLEDGMENTS
We thank Dr. J. M. Trappe for preparing the Latin diagnoses, reviewing the manuscript and making useful
suggestions, Dr. J. Morton for providing spores and helpful comments, R. Sheath, A. Steinman, J. Moffet
and C. Friese for collecting dune samples, D. Scales for the scanning electron microscopy, US National
Park Service for permission to collect samples from National Park lands, the Chief Ranger of Seashore
State Park, and Don Tiller, Refuge Manager, US Fish and Wildlife Service, for permission to collect at the
Moonstone Beach dunes, Rhode Island, W. L. Halvorson, Channel Islands National Park for making
possible the collections from California, T. H. Nicolson for his advice, and Marion Jenkins and
Shiela Swan for typing the manuscript.
255
LITERATURE CITED
Anon, 1969. Flora of the British Fungi. Colour Identification Chart. Royal Botanic Garden, Edinburgh.
Ferrer, R. L., and R. A. Herrera, 1981. El genero Gigaspora Gerdemann et Trappe (Endogonaceae) en
Cuba. Rev. Jardin Bot. Nacional, Habana 1(1):43-66.
Hall, I. R., 1984. Taxonomy of VA mycorrhizal fungi. In: VA Mycorrhiza (Ed. by C. L. Powel &
D. J. Bagyaraj), pp. 57-95, CRC Press, Florida.
Hall, I. R. and L. K. Abbott, 1984. Some Endogonaceae from South Western Australia. Transactions of
the British Mycological Society 83:203-208.
Koske, R. E., and B. Tessier, 1983. A convenient, permanent slide mounting medium. Mycological Society
of America Newsletter 34(2):59.
Koske, R. E., and C. Walker, 1984. Gigaspora erythropa, a new species forming arbuscular mycorrhizae.
Mycologia 76: 250-255.
Morton, J. B., 1986a. Three new species of Acaulospora from high aluminum, low pH soils in West
Virginia. Mycologia 78. In press.
Morton, J. B., 1986b. Effect of mountants and fixatives on wall structure and Melzer’s reaction in spores of
two Acaulospora species (Endogonaceae). Mycologia. In review.
Nicolson, T. H. and Schenck, N. C., 1979. Endogonaceous mycorrhizal endophytes in Florida. Mycologia
71:178-198.
Walker, C., 1979. The mycorrhizast and the herbarium: the preservation of specimens from VA
mycorrhizal studies. In: Program and Abstracts, 4th N. American Conference on Mycorrhiza.
Fort Collins, Colorado.
Walker, C., 1983. Taxonomic concepts in the Endogonaceae: spore wall characteristics in species
descriptions. Mycotaxon 18:443-455.
Walker, C., 1986. Taxonomic concepts in the Endogonaceae: II. A fifth morphological wall type in
endogonaceous spores. Mycotaxon 25:95-99.
Walker, C. and Sanders, F. E., 1986. Taxonomic concepts in the Endogonaceae: III. The separation of
Scutellospora gen. nov. from Gigaspora Gerd. & Trappe. Mycotaxon. In Press.
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MYCOTAXON
Vol. XXVII, pp. 237-245 October-December 1986
FOUR NEW EFFIGURATE-CRUSTOSE SPECIES OF PARMELIA
(LICHENES, PARMELIACEAE) FROM SOUTHERN AFRICA
FRANKLIN A. BRUSSE
Botanical Research Institute
Private Bag X101, Pretoria,
South Africa
ABSTRACT
Four new effigurate-crustose species are described and one new combination is made
in the lichen genus Parmelia from southern Africa. The new species are Parmelia insipida
Brusse, P. nimbicola Brusse, P. ralla Brusse and P. squamatica Brusse. The new combina-
tion is Parmelia leptoplaca (Zahlbr.) Brusse. The affinities, chemistries and distributions
of these species are discussed.
INTRODUCTION
The five species dealt with here, are rather unusual for the genus Parmelia in that they
have crustose thalli with effigurate margins. The lower cortex is fused to the rock sub-
strate, except at the lobe apices, where the free lower surface can be observed. True rhi-
zines are not present. In addition, the apothecia are much smaller than is usual for Par-
melia, and sometimes remain immersed in the central areoles. This variation does not
appear to be significant for a similar case exists in the genus Tephromela, where the
apothecia may be large (and resemble those of the large majority of Parmeliae) or much
smaller and immersed in areoles. The internal structure of the apothecia of these new
species is consistent with that of Parmelia, except for a somewhat thicker subhymenium
in the central area of the apothecium.
The new species cannot be adequately accomodated in either of the effigurate-crustose
genera, Protoparmelia Choisy or Protoparmeliopsis Choisy. Protoparmelia, typified by
P. badia (Hoffm.) Choisy, seems to be quite distinct because of the well separated, capped
paraphyses, elongated ascospores (with length more than twice the breadth, and a more
strongly amyloid tholus (Hafellner, 1984). Here the pycnidiospores are acicular as in
Parmelia. On the other hand Protoparmeliopsis, typified by Protoparmeliopsis muralis
(Schreb.) Choisy, cannot be used for these lichens, because there the pycnidiospores are
long and thread-like, commonly being over 20 um long and perhaps more importantly,
being only around 0.5 um in diameter (Poelt, 1958). However, the ascus of Protoparme-
liopsis is poricidal with extension (as with Parmelia) and the tholus reacts in a similar
manner to that of Parmelia in Lugols iodine solution.
238
NEW SPECIES
PARMELIA INSIPIDA Brusse, sp. nov.
Thallus crustosus et effiguratus, saxicola, usque ad 4 cm diametro. Lobi elongati,
0.25—1.0 um lati, 830—130 um crassi. Thallus superne viridis, emaculatus, isidiis sorediis-
que nullis. Cortex superior 20 ym crassus. Stratum gonidiale 30—65 ym crassum, algis
Trebouxiis, 6—20 um diametris. Medulla alba, 25—65 um crassa. Cortex inferior 18 um
crassus, ad apices loborum solum bene evolutus. Rhizinae non bene evolutae. Apothecia
immersa vel leviter elevata, usque ad 0.7 mm diametris. Hypothecium 28—33 um cras-
sum, Subhymenium 10—60 yum crassum. Hymenium 50—65 ym altum, J+ caeruleum.
Asci clavati, cum tholis J+ caeruleis (fig. 1). Ascosporae octonae, ovales, hyalinae, 6.5—
10.5 x 5.0—7.5 um. Pycnidia globosa, 80—110 x 50—90 um. Pycnidiosporae bacillares,
6.0—11.5 x 0.8 um. Thallus acidum usnicum et acidum evernicum continens.
TYPUS: Cape, 3124 (Hanover): Lootsberg Pass between Graaff-Reinet and Middel-
burg, Sneeuwberg, on siltstone kranz and boulders on NE slope, alt. 1 800 m, (-DD)
F. Brusse 4692, 1986.01.30 (PRE, holo-; BM, LD, US, iso-), fig. 4.
Thallus crustose and effigurate, saxicolous, up to 4 cm across. Lobes elongate, 0.25—
1.0 mm broad, 80—130 um thick. Upper surface green, lacking maculae, isidia and sore-
dia. Upper cortex 20 wm thick. Algal layer 30—65 um thick, algae Trebouxia, 6—20 um
diam. Medulla white, 25—65 um thick. Lower cortex 18 yum thick, well developed only
near the lobe tips. Rhizines not well developed. Apothecia immersed or only slightly
emergent, up to 0.7 mm across. Hypothecium 28—33 um thick. Subhymenium 10—60
yum thick. Hymenium 50—65 um high, J+ blue. Asci clavate, eight-spored, tholus J+ blue
(fig. 1). Ascospores oval, hyaline, 6.5—10.5 x 5.0—7.5 um. Pycnidia globose, 80—110 x
50—90 yum. Pycnidiospores rods, 6.0—11.5 x 0.8 um. Chemistry: Usnic acid in the up-
per cortex and evernic acid in the medulla.
This species resembles the more common Parmelia leptoplaca (Zahlbr.) Brusse, with
which it sometimes grows on the same rock. The latter species is a little larger and deeper
yellow in colour, and can be separated from this new species by the presence of salazinic
acid in the medulla.
Parmelia insipida is known from the eastern Cape Province, from the Sneeuwberg
north of Graaff-Reinet to the south-western end of the Drakensberg north of Elliot.
Parmelia leptoplaca is much more widespread and common, being present throughout
the Great Karoo in a broad sense.
CAPE.—3127 (Lady Frere): 17 km N of Elliot, Barkleys Pass, on Clarens sandstone
kranz on SW slope, alt. 2050 m (—BB), F. Brusse 4616, 1986.01.27 (PRE).
PARMELIA NIMBICOLA Brusse, sp. nov.
Thallus crustosus et effiguratus, saxicola, usque ad 1.5 cm diametro. Lobi elongati,
0.2—0.6 mm lati, 100—115 um crassi. Thallus superne cinereus, emaculatus, isidiis sore-
diisque nullis. Cortex superior 15—20 wm crassus. Stratum gonidiale 30—45 um crassum,
algis Trebouxiis, 4-17 mm diametris. Medulla alba, 50—60 um crassa. Cortex inferior
6—10 pm crassus, ad apices loborum solum bene evolutus. Thallus inferne pallidus. Rhi-
zinae non bene evolutae. Apothecia sessilia, usque ad 1 mm diametris. Hypothecium
25—45 um crassum. Subhymenium 20—50 wm crassum. Hymenium 45—55 um altum,
239
Fig. 1. Parmelia insipida Brusse, ascus and paraphyses. F. Brusse 4692, holotype. Bar =10 pum.
J+ caeruleum. Asci clavati, cum tholis J+ caeruleis (fig. 2). Ascosporae octonae, ellip-
soideae, 8.0—11.5 x 5.5—7.0 um. Pycnidia ignota. Thallus atranorinum et acidum suc-
cinoprotocetraricum continens.
TYPUS: Cape, 3322 (Oudtshoorn): Robinsons Pass between Mossel Bay and Oudts-
hoorn, SW slopes of Ruitersberg, on Table Mountain sandstone outcrops, alt. 790 m
(—CC), F. Brusse 4807, 1986.02.04 (PRE, holo-; BM, LD, iso-), fig. 5.
Thallus crustose and effigurate, saxicolous, up to 1.5 cm across. Lobes elongate, 0.2—
0.6 mm broad, 100—115 um thick. Upper surface grey, lacking maculae, isidia and sore-
dia. Upper cortex 15—20 um thick. Algal layer 30—45 mm thick, algae Trebouxia, 4—17
gum diam. Medulla white, 50—60 um thick. Lower cortex 6—10 um thick. Lower surface
pale. Rhizines not well developed. Apothecia sessile, up to 1 mm across. Hypothecium
25—45 ym thick. Subhymenium 20—50 um thick. Hymenium 45—55 um high, J+ blue.
Asci eight spored, clavate, tholus J+ blue (fig. 2). Ascospores ellipsoid, 8.0—11.5 x 5.5—
7.0 um. Pycnidia unknown. Chemistry: Atranorin in the upper cortex and succinopro-
tocetraric acid in the medulla.
240
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Fig. 2. Parmelia nimbicola Brusse, ascus and paraphyses. F. Brusse 4807, holotype. Bar = 10 pm.
This new species is the first crustose Parmelia with atranorin in the upper cortex. The
recently described P. astricta (Brusse, 1984), P. fausta, P. insignis and P. pudens (Brusse,
1986), are also narrow lobed small species, but they are clearly foliose and not crustose.
P. nimbicola was found on the western end of the Outeniqua mountains, north of Mos-
sel Bay. This is an unusual habitat for a crustose Parmelia since this locality is very often
shrouded in mist and the rainfall is relatively high. There are no close relatives of this
species known at present.
P. nimbicola is presently known only from the type locality.
PARMELIA RALLA Brusse, sp. nov.
Thallus crustosus et effiguratus, interdum dispersus, saxicola, usque ad 3.5 cm dia-
metro. Lobi elongati vel sublineares, 0.2—0.5 mm lati, 100—120 wm crassi. Thallus su-
perne viridis, emaculatus, sorediis isidiisque nullis. Cortex superior 10—20 um crassus.
Stratum gonidiale 30—70 wm crassum, algis Trebouxiis, 5—18 um diametris. Medulla
alba, 15—50 um crassa. Cortex inferior 9—13 ym crassus, ad apices loborum solum bene
241
Fig. 3. Parmelia ralla Brusse, ascus and paraphyses. F. Brusse 4935, holotype, Bar =10 gan.
evolutus. Rhizinae non bene evolutae. Apothecia sessilia, usque ad 0.7 mm diametris.
Hypothecium 30—55 wm crassum. Subhymenium 25—40 ym crassum. Hymenium 45—
55 um altum, J+ caeruleum. Asci clavati, tholis J+ caeruleis (fig. 3). Ascosporae octonae,
ellipsoideae, 8.0—10.5 x 5.0—7.0 um. Pycnidia globosa, 100—150 x 85—120 um. Pycni-
diosporae bacillares, 7.5—13 x 1 um. Thallus acidum usnicum et norlobaridonum conti-
nens.
TYPUS: Cape, 3321 (Ladismith): Swartberg range, Seven Weeks poort, 8,6 km NW
of Amalienstein, on Table Mountain sandstone boulders on SW slope, alt. 670 m (—AD),
F. Brusse 4935, 1986.02.10 (PRE, holo-; BM, COLO, LD, MEL, US, iso-), fig. 6.
Thallus crustose and effigurate, sometimes dispersed, saxicolous, up to 3.5 cm across.
Lobes elongate to sublinear, 0.2—0.5 mm broad, 100—120 um thick. Upper surface
green, lacking maculae, isidia and soredia. Upper cortex 10—20 um thick. Algal layer
30—70 um thick, algae Trebouxia, 5—18 um diam. Medulla white, 15—50 um thick.
Lower cortex 9—13 ym thick, well developed at lobe apex only. Rhizines not well de-
242
veloped. Apothecia sessile, up to 0.7 mm across. Hypothecium 30—55 um thick. Sub-
hymenium 25—40 ym thick. Hymenium 45—55 yum high, J+ blue. Asci clavate, eight-
spored, tholus J+ blue (fig. 3). Ascospores ellipsoid, 8—10.5 x 5.0—7.0 um. Pycnidia
globose, 100—150 x 85—120 um. Pycnidiospores rods, 7.5—13 x 1 yum. Chemistry:
Usnic acid in the upper cortex and norlobaridone in the medulla.
This new species resembles other crustose Xanthoparmeliae, but has more elongated,
sublinear lobes. The norlobaridone in the medulla readily distinguishes this new species
from morphologically similar taxa. P. ralla has been found from Uniondale to Ladismith,
on Table Mountain sandstone.
Cape.—3323 (Willowmore): 5 km § of Uniondale, Uniondale poort, on steep SW
slope, on TMS outcrops, alt. 830 m (—CA), F. Brusse 4768, 1986.02.03 (PRE).
PARMELIA SQUAMATICA Brusse, sp. nov.
Thallus crustosus et effiguratus, saxicola, usque ad 5 cm diametro. Lobi elongati,
0.2—0.7 mm lati, 100—300 mum crassi. Thallus superne viridis, emaculatus, sorediis isi-
diisque nullis. Cortex superior 10—20 um crassus. Stratum gonidiale 30—80 ym crassum.
Medulla alba, 60—200 wm crassa. Cortex inferior 10O—15 um crassus, ad apices loborum
solum bene evolutus. Thallus inferne pallidus. Rhizinae non bene evolutae. Apothecia
non visa. Pycnidia circa 50 um diametris, globosa. Pycnidiosporae bacillares, 7—11 x 1
um. Thallus acidum usnicum et acidum squamaticum continens.
TYPUS: Cape, 3219 (Wuppertal): Summit of Pakhuis Pass, near Clanwilliam, on
TMS boulders and outcrops on SW slope, alt. 900 m (—AA), F. Brusse 772 8-2-1,
1977.02.08 (J, holo-), fig. 7.
Thallus crustose and effigurate, saxicolous, up to 5 cm across. Lobes elongate, 0.2—
0.7 mm broad, 100—300 um thick. Upper surface green, lacking maculae, isidia and
soredia. Upper cortex 10—20 um thick. Algal layer 30—80 um thick. Medulla white,
strongly UV366 fluorescent (white), 60—200 um thick. Lower cortex 10—15 um thick,
well developed at lobe tips only. Lower surface pale, without well developed rhizines.
Apothecia not seen. Pycnidia around 50 yum diam., globose. Pycnidiospores rods, 7—11
x 1 um. Chemistry: Usnic acid in the upper cortex and squamatic in the medulla.
This new species resembles other crustose Xanthoparmeliae, but can be distinguished
from them most readily by the presence of squamatic acid in the medulla, which causes
a strong white fluorescence in long wave ultraviolet light.
At present this species is known only from the type locality.
NEW COMBINATION
PARMELIA LEPTOPLACA (Zahlbr.) Brusse, comb. nov.
Basionym: Lecanora leptoplaca Zahlbruckner, A., Ann. Crypt. Exot. 5: 249, 1932.
TYPUS: Kapland, Laingsburg (grosse Karroo), Sandsteinfelsen, c. 700 m, J. Brunnt-
haler, 11.11.1909 (W 626!).
243
Fig. 4. Parmelia insipida Brusse, habit F. Brusse 4692, holotype. Scale in mm and cm.
Fig. 5. Parmelia nimbicola Brusse, habit. F. Brusse 4807, holotype. Scale in mm and cm.
244
Fig. 6. Parmelia ralla Brusse, habit. F. Brusse 4935, holotype. Scale in mm and cm.
Fig. 7. Parmelia squamatica Brusse, habit. F. Brusse 772 8—2—1, holotype. Scale in mm.
245
ACKNOWLEDGEMENTS
The author is grateful to the South African Department of Agriculture and Water
Supply for financial support. Thanks are also extended to Dr John A. Elix for reviewing
this paper. The author is also thankful to Mrs A.J. Romanowski for the photography
and to Mrs S.S. Brink and Mrs E. Bunton for the typing.
LITERATURE CITED
BRUSSE, F.A. 1984. New species and combinations in Parmelia (Lichenes) from south-
ern Africa. Bothalia 15 (1 & 2): 315-321.
BRUSSE, F.A. 1986. Five new species of Parmelia (Lichenes, Parmeliaceae) from south-
ern Africa. Mycotaxon in press.
HAFELLNER, J. 1984. Studien in Richtung einer nattirlichen Gliederung der Sammel-
familien Lecanoraceae und Lecideaceae. Beih. Nova Hedwigia 79: 241—371.
POELT, J. 1958. Die lobaten Arten der Flechtengattung Lecanora Ach. sensu ampl. in
der Holarktis. Mitt. bot. Staatss. Mtinchen 2: 411—573.
MYCOTAXON
VOISIXAVIL S ppe 247-2535 October-December 1986
BERTIELLA (SACC.) SACC. & SYDOW
A SYNONYM OF MASSARINA SACC.
Ove Eriksson
Institute of Ecological Botany,
University of Umea, S-90187 Umea, Sweden.
and
Jing-zhu Yue
Institute of Microbiology,
Academia Sinica, Beijing, PRC - China.
ABSTRACT
The generic type Bertiella macrospora (Sacc. )
Sace.& Sydow is a Massarina species, M.
macrospora (Sacc.) 0. Eriksson & J.-z. Yue.
It is compared with some Lophiostoma spp. and
M. coccodes (Karsten) 0. Eriksson & J.-z.
Yue.
INTRODUCTION
During our studies of Chinese pyrenomycetes, we examined
material identified as Bertia macrospora Sacc. (Tai 1979:
89, Herb. HMAS) and found that all collections were Bertia
moriformis (Tode: Fr.) De Not. var. latispora Corlett &
Krug. However, Nannfeldt (1975: 291) reported that B.
macrospora, type of the genus Bertiella (Sacc.) Sacc. &
Sydow, is a bitunicate fungus. In order to clarify the true
systematic position of this genus, we examined original
material of B. macrospora. We found that Nannfeldt's
Opinion is correct, and that the species should be referred
to Massarina Sacc.
MATERIALS AND METHODS
Methods used in this study are described in Eriksson & Yue
(1985) and Yue & Eriksson (1985).
248
TYPE OF BERTIELLA (SACC.) SACC.
Massarina macrospora (Sacc.) 0. Eriksson & J.-z. Yue, comb.
nov. - Basionym: Bertia macrospora Sacc., Michelia 1:
452 (1878).
Pig oe.
Ascomata superficial on wood, but possibly originally
covered by cortex, - gregarious and sometimes confluent,
subglobose with a flat base or more irregular in shape;
surface often slightly striated along the grain, non-papil-
late, non-setose, black, carbonaceous. Ascomal wall ca.
60-70 ym thick, partly consisting of ~ isodiametric cells
with strongly melanized, dark brown walls in outer parts of
ascomata, with thin-walled hyaline walls in inner parts,
partly of less dark-coloured bands of elongated cells,
somewhat resembling a cephelothecioid cell arrangement.
Hamathecium of numerous, filiform, ca. 2 ym wide, branched
paraphysoids. Asci numerous, cylindric-clavate, short-
stalked, ca. 130-150 x 22 pm, bitunicate. Ectotunica thin;
endotunica very thin in lower half of mature ascus, thicker
Fig. 1. Massarina macrospora. (A). Asco-
mata. (B). Ascus. (C). Ascospores, hya-
line, 1-septate. (D). Ascospores?, brown,
3-septate. - Coll.: Sace., Mycoth. Ven.
651 (UPS). -— Magn.: (A) x22. (B-D) scale.
249
towards the apex and with a small ocular chamber, but no
ring structures, its innermost parts staining dark with
Chlorazol black. Ascospores ca. 37-43 x 8-9 um, fusiform,
inequilateral, 1-septate, hyaline, with 2-3 globules in
each hemispore, smooth, without gelatinous sheath.
On wood of Fagus sylvatica, Italy, Treviso, Consiglio, P.A.
Saccardo, Mycoth. Ven. 651 (UPS).
DISCUSSION
Bertia subg. Bertiella Sacc. (1882: 584) was raised to
generic rank by Saccardo (1899: 19). The subgenus original-
ly contained two species, Bertia macrospora Sacc. and B.
parasitica Fabre. Clements & Shear (1931: 270) selected B.
macrospora as type species of the genus Bertiella (Sacc.)
Sace. & Sydow. It should be mentioned that Kirschstein
(1906: 51) used the name Bertiella for another genus, based
0 100 pm
Fig. 2. (A-C). Lophiostoma subcorticale. (D, E). L. semi-
liberum. (F). L. arundinis. - (A, D). Ascomata. (B, C, E,
F). Ascospores. - Coll.: (A-C). K. & L. Holm 3415a (UPS).
(DE). O. Eriksson 2376d (UME 25621). (F). 0. Eriksson 2490b
COPS). = Magne: CA, 3D) ex 22. (BSC, BF) seales
250
on Bertiella polyspora Kirschst. = Kirschsteinia polyspora
(Kirschst.) H. Sydow (1906: 455), which was subsequently
(Kirschstein 1912: 289) identified as Fracchiaea hetero-
gena Sacc. This name is a synonym of Nitschkia broomeiana
(Berk.) Nannf.
The type species of Bertiella (Sacc.) Sacc. & Sydow,
Bertia macrospora, was described by Saccardo in 1878. We
studied original material (see above) and found that this
species belongs to Massarina. This genus is characterized
by perithecioid ascomata which are often almost hyaline in
basal parts and unevenly pigmented in upper parts (opaque
near the pseudostiole). They are immersed in most species,
e.g. the type M. eburnea (Tul. & C. Tul.) Sacc., but some
species have erumpent ascomata. The centrum consists of
paraphysoids and bitunicate asci. These are rather thin-
walled and produce 1-5-septate, hyaline ascospores. In some
species the spores lack a distinct perispore, in others
they are completely enclosed in a gelatinous layer. In one
species, a potential pathogen of man in the Tropics, which
is currently known as Leptosphaeria senegalensis Segretain
& al. (not validly published, ICBN Art. 37, which is why
transfer to Massarina must await), the ascospores have a
thick perispore with a characteristic appendage at the
lower end (El-Ani & Gordon 1965: Figs. 1-4; Parguey-Leduc &
al. 1982: Figs. 6-7). Massarina macrospora deviates from
the majority of the species in the genus in their ascomata
being almost superficial when mature.
We checked the reference literature for older names
which might possibly be synonyms of B. macrospora, but none
of the species in Didymella D. Sacc., Massarina, Meta-
sphaeria Sacc., Sphaeria, or other relevant genera could
possibly be conspecific with B. macrospora. However, Massa-
rina is close to Lophiostoma Ces. & De Not. (incl. Lophio-
trema Sacc.), the compressed ostiolar neck in this genus
being the main distinguishing character from Massarina, As
normal specimens with compressed ostiole are sometimes
intermixed with others without such a neck, there was the
possibility that the type collection of B. macrospora is
only an atypical Lophiostoma species without compressed
neck. Therefore, we also searched for possible synonyms in
Lophiosphaera Trev., Lophiotrema, and Lophiostoma. We found
that three species with older epithets may have similar
hyaline, 1-septate ascospores.
1. Lophiostoma subcorticale Fuckel (Fig. 2 A-C) has
spores with much the same shape as spores of B. macrospora,
but they are normally about 50 % longer and wider (ca. 60 x
12 ym versus ca. 40 x 8 pm). Chesters & Bell (1970: 12)
co
have given spore measurements of 45-60 x 8-10 pm for L.
subcorticale, but this interval probably includes measures
for very young, immature spores. Fully ripe spores of L.
subcorticale are multi-septate and brown. In one slide of
B. macrospora we saw 3-septate, brown spores near ascomata,
but we are not sure that they belong to that species. The
ascomata of L. subcorticale are usually immersed in bark
and the protruding necks distinctly compressed. Ascomata of
M. macrospora are superficial on naked wood, but may
originally have been covered by cortex. They have no necks.
2. Lophiostoma angustilabrum (Berk. & Br.) Cooke has
shorter and narrower spores (cf. Chesters & Bell 1970: 8).
3. L. semiliberum (Desm.) Ces. & De Not.) (Fig. 2 D-F)
is a graminicolous species with hyaline, 1-septate spores
measuring about 34-38 x 5-6 pm. The ascomata are often
more or less erumpent and the necks may be compressed,
non-compressed or missing, but there always seems to be
some ascomata with compressed neck in the same sample.
Lophiostoma arundinis (Pers.: Fr.) Ces. & De Not., a
similar species with 5-septate, brown spores, has been
considered the mature stage of L. semiliberunm.
Our conclusion is that there is no older epithet for
B. macrospora, and that the correct name for the species is
Massarina macrospora (Sacc.) 0. Eriksson & J.-z. Yue.
In the dissecting microscope M. macrospora resembles
Leptosphaeria coccodes Karsten, but the ascomal wall is
thinner and of more homogeneous texture in that species,
and the spores are distinctly swollen above the primary
septum (see Eriksson 1967: 401). Munk (1956: 303) treated
L. coccodes as the type of Metasphaeria Sacc. in the new
family Massarinaceae. However, Clements & Shear had already
cited "M. sepincola (Fr.) Sacc!" as type for Metasphaeria in
1931 (this type was accepted by Holm in Farr & al. 1979:
1078). This is now called Saccothecium sepincola (Fr.) Fr.,
a species which is not related to Metasphaeria sensu Munk.
There are, however, no important differences between the
latter taxon and Massarina, and the correct name for Kars-
ten’s fungus is Massarina coccodes (Karsten) 0. Eriksson &
J.-z. Yue comb. nov. (bas. Leptosphaeria coccodes Karsten,
Fungi Fenn. exs., no. 963, 1870).
As mentioned above, the only difference between Massa-
rina and Lophiostoma is the presence of compressed ostiolar
necks in normal specimens of the latter genus. This is
probably not a very important difference, but, for the time
being, we suggest that the genera are kept separate, and
that species with erumpent ascomata, lacking a compressed
ostiolar neck, are placed in Massarina. However, future
252.
studies of the whole group may result in redefined generic
limits. In any case, presence of compressed ostiolar neck
is not a criterion of importance at the familial level, and
the family Massarinaceae should be included in the Lophio-
stomataceae.
Lophiosphaera beckhausii (Nits.) Berl. & Vogl. may be
a younger synonym for Massarina macrospora, but we have not
seen any original material. The species is not treated by
Chesters & Bell (1970).
ACKNOWLEDGEMENTS
We acknowledge with thanks the financial support for our
work from Academia Sinica (Beijing) and the Swedish Natural
Science Research Council (Stockholm). We are grateful to
Prof. Dr. N. Lundquist, Dr. L. Holm and Dr. V. Shingler for
reviewing the manuscript.
LITERATURE CITED
BOSE, S.K., 1961. Studies on Massarina Sace. and related
genera. - Phytopath. Z. 41: 151-213.
CHESTERS, C.G.C. & BELL, A.E., 1970. Studies in the Lophio-
stomataceae Sacec. - Mycol. Pap. 120: 1-55.
CLEMENTS, F.E. & SHEAR, S.L. 1931. The Genera of Fungi. -
Hafner, New York.
EL-ANI, A.S. & GORDON, M.A. 1965. The ascospore sheath and
taxonomy of Leptosphaeria senegalensis. - Mycologia 57:
275-278.
ERIKSSON, 0O. 1967. On graminicolous pyrenomycetes from
Fennoscandia. - Ark. Bot. 6: 381-440.
- & YUE, J.-Z. 1985. Studies on Chinese ascomycetes.
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SYDOW, H. 1906. Referate und kritische Besprechungen - Ann.
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VOWe Oey es, Pps <2o ome OL October-December 1986
A NEW CHLAMYDOMYCES SPECIES FROM ITALY
PERINI CLAUDIA
Dipartimento di Biologia Ambientale - Sezione Botanica
Universita di Siena - Via P.A. Mattioli 4, 53100 Siena, Italy
Abstract
A new species of the monospecific genus Chlamydomyces, C. sympodialis,
found on dead leaves of Quercus pubescens, is here described and
illustrated.
An interesting Hyphomycete on dead leaves of Quercus pubescens
was found in the Botanical Garden of Rome in April 1985. It produces
1-septate conidia and possesses an Aspergillus-like synanamorph.
The following description concerns observations on natural media;
the attempt at isolation in pure culture (V-8 Agar) was successful.
Exsiccatum is available in the Mycological Herbarium of the
Rome Botanical Garden (ROHB 131 C).
Chlamydomyces sympodialis sp. nov.
Coloniae effusae, floccosae, hyalinae, conidiis pallide inauratae-
-brunnae, mycelium conidiaque valde refracta. Mycelium superficiale.
Conidiophora oriunda sicut ramificationes ex hyphis' reptantibus
vel aereis, macronematosa, mononematosa, flexuosa, geniculata,
simplicia interdum rectis angulis ramosa, hyalina, septata, laevia,
usque ad 325 pum longa, 2.5-5 pm lata, rami usque ad 185 pm longi.
Cellulae conidiogenae monoblasticae vel polyblasticae, integratae,
terminales vel intercalares, denticulatae, 80-130 pm longae; denticu-
lus. subconicus,. in basi septatus, et in apice apertus, 2.3-5 ym
longus, 1.5-2.7 pm in basi latus. Conidia solitaria, acropleurogena,
1-septata, napiformi-tubrinata, laevia, 26.5-39 x 18.5-25.5 pam;
cellula superior subglobosa et in basi trunca, inaurata—brunnea,
14-23.5 pm; cellula inferior obconica fere, evadens in brevem apertam
et truncam. appendicem, pallida inaurata-brunnea. Status Aspergillus-—
-Similis: conidiophora mononematosa, simplicia, hyalina, septata,
usque ad 56.5 pm longa; vesiculae subglobosae; phialides uniseriatae,
ampulliformes, 6.5-10 x 3.5-6 pm. Phialoconidia catenulata, ellipsoi-
dea vel subglobosa, laevia, 3.3-6 x 2.5-3.3 pm.
256
In foliis emortuis Q. pubescens, Rome, holotypus: ROHB 131 C.
Colonies effuse, floccose, hyaline, pale golden-brown after
sporulation, mycelium and conidia refracting. Mycelium superficial.
Conidiophores arising as branches from aerial or reptant vegetative
hyphe, macronematous, mononematous, flexuous, geniculate, simple
or sometimes branched at right-angles, septate, smooth, hyaline,
up to 325 pm long, 2.5-5 pm wide, branches up to 185 pm long. Conidio-
genous cells polyblastic or monoblastic, integrated, terminal
or intercalary, denticulate, 80-130 pm long; denticles subconical,
septate. at the base and opened at the apex, 2.3-5 pm long .and)1.5-
-—2.7 pm wide at the base. Conidia solitary, acropleurogenous,1-septate
turbinate-napiform, smooth, 26.5-39 x 18.5-25.5 um; distal cell
subglobose and truncate at the base, golden-brown, 14-23.5 pm long;
proximal cell more or less obconical, terminating in a Short, opened
and truncate appendage, pale golden-brown. Aspergillus-like state:
conidiophores consisting of a mononematous, Simple, hyaline, septate
stipe, up to 56.5 pm long and 3.3-5 pm wide, usually terminating
in, 'a ‘subglobose. vescicle, 11.5-15 x 0-12.51 jm: Phialides sites
single series, ampulliform, 6.5-10 x 3.5-6 jm, produced numerous
on the vescicle; phialides are sometimes produced on the unswollen
apex of the stipe. Phialoconidia catenulate, subglobose to ellipsoid,
hyaline, smooth, 3.3-6 x 2.5-3.3 pm.
The new species described is characterized by the presence of
two kind of conidia. The small, O-septate phialoconidia are produced
in an Aspergillus-like stage, while the holoblastic ones are produced
in a stage characterized by the presence of denticulate conidiophore
and conidiogenous cells with a sympodial-like stucture.
Holoblastic conidiogenesis takes place with the swelling of
the hyphal apex which, even at this early stage, greatly resembles
the conidium. Below the slight constriction between the growing
conidium and the conidiogenous cell apex develops a septum that,
after the conidial secession, will identify with the basal septum
of the denticle. Subsequently, a second septum develops in the
submedian area of the conidium. On reaching maturity it detaches
itself by means of a rhexolitic rupture in an especially weak point
in the wall. Separation occurs in the lower part of the cell delimited
by the basal septum of the denticle and by the submedian septum
of the conidium. This cell, which presents cytoplasmatic continuity,
can, at this stage, be considered a separating cell. After rupture,
the basal part of the latter cell on the conidiogenous cell is
identifiable as a subconical denticle septate at the base and open
at the apex. The other portion, of a larger dimension, will become
the proximal cell of the conidium that appears closed by a cytoplasma-—
tic accumulation slightly above rupture point.
The conidia have been described as bicellular because, when
A) conidiophore and conidia
B) Aspergillus-like synanamorph
258
Fig. 2 -— Chlamydomyces sympodialis sp. nov.
a) conidia
b) conidiophores
c) denticles
d) Aspergillus-like synanamorph
259
detached, they are formed by two cells, only the upper one of which
is probably the real spora. Although the base of the proximal obconi-
cal cell is undelimited by a wall structure to divide it from the
appendix,it conserves its cytoplasmatic content even after separation.
For this reason it is considered as belonging, in every respect,
to the separated conidium. In fact, separation is considered the
final stage of a conidiogenetic development and, for this reason,
the mature conidium should therefore be described after, and not
before, the actual separation.
The aspect of the conidiophore presumes a typically sympodial
development.
In 1907 Bainier describes the new genus Chlamydomyces in the
following way: ''...Conidies bicellulaires a 1l'extrémité de supports
a plusieurs étages de ramifications généralment opposées. Cellule
supérieure sphérique ou ovale, tronquée a sa base, a paroi épaisse
et de couleur accentuée. Cellule inférieure en tronc de céne renversé
a paroi mince, lisse, incolore et translucide.'"' No original material
remains of the type species C. diffusus, that, in according to
Bainier (1907), '"...posséde un mycélium et des appareils fructiféres
d' Acremoniella_atra, donnant naissance a des spores de Mycogone
cervina.'' Mason (1928) however, accepts the new genus Chlamydomyces
for species that present "...large red or yellow two-celled macrospo-
res, with maximum dimensions of about 40 x 28 pm" and that differ
from Mycogone "... by the small, broadly cuneate (not subglobose)
hyaline, smooth basal cell and by the fact that there is no con-
striction at the septum" and that, further, "...form an Aspergillus-—
-like microconidial stage in culture." In this occasion the Author
publishes the new combination C. palmarum rendering synonymous
Trichobasis palmarum Cooke (1877), Uredo palmarum (Cooke) De Toni
(1888), Mycogone rufa Petch, 1917 and proposing that C. diffusus,
the type species of the genus, might also be considered among the
synonyms. Finally, in 1976, Ellis nominates C. diffusus as a synonym
of C. palmarum.
For a better understanding of the studied species the following
authentic material has been examined:
Chlamydomyces palmarum (Cooke) Mason, MFC - 9083 - Host: Curcuma
aromatica leaves; local.: Kuantzuling, Taiwan; Date: 7/11/1978
Ref.: Mats. Myc. Mem. 1:17, 1980
Mycogone rufa Petch, Herb. Hort. Reg. Bot. Kew. type - Host: plaintain
fruit; Peradenija 4542; Date: 16/3/1915
Trichobasis palmarum Cooke, Herb. Hort. Reg. Bot. Kew, type - Host:
cocopalm leaves; Demerara -— Uredo 135 3084
The latter specimen was found to be quite old which rendered
260
study very difficult. (Even when the species has been identified,
it is extremely difficult to interpret particular structures).
The C. palmarum conidiophores are either simple or branched,
branching being variable both in number and symmetry, often at
right angles and sometimes composed of alternate placings. The
conidia are bicellular; the upper cell is echinulate and thick-walled
whereas the thin-walled, proximal obconical cell is sometimes peduncu-
late.
During holoblastic conidiogenesis two septa are subsequently
formed: the first delimits the young, swollen conidium from the
underlying cell; the second, being formed in the submedian position,
divides the conidium in two cells. The underlying cell of the first
septum formed appears to be divided by a stucture of difficult
identification. According to Howell (1939) it seems a wall structure;
but a careful microscopic study identicates that it is simply an
area of a different cytoplasmatic density. Such a cell, considered
conidiogenous, is lacerated during rhexolitic separation. Its apical
portion sometimes remains as a peduncle at the base of the conidiun;
contrarily, the basal portion, of larger dimensions, remains on
the conidiophore as a denticle, septate at the base and delimited
apically by a cytoplasmic structure.
The presence of a conidiogenetic process and of a _ rhexolitic
secession identical in all phases to those of C. palmarum and also
of bicellular conidia and an Aspergillus-like stage, would seem
to allow the inclusion of the species in Chlamydomyces. The species
described differs, however, from the type species by its geniculated
structure of the conidiophore and by the smooth apical cells of
the conidia. The morphology of the conidiophore is less characteristic
in C. palmarum and the conidia have a thick-walled, echinulate
upper cell and a smooth, thin-walled basal cell. The form and dimen-
sions of the Aspergillus-like stage also differ.
For the above mentioned reasons the new species C. sympodialis
is proposed.
Acknowledgements
The Author whish to thank Prof. A. Rambelli and Dr. S. Onofri,
Cattedra di Micologia, Dipartimento di Biologia Vegetale, Universita
degli Studi di Roma "La Sapienza", Rome, Italy, for their helpful
suggestions and for kindly reviewing the manuscript.
REFERENCES
Bainier, M.G. 1907. Mycothéque de 1'Ecole de Pharmacie de Paris.
Bull. trim. Soc. Mycol. Fr.',ni23$235-241y. pls 26
Cooke, M.C. 1877. Cocoa-Palm fungi. Grevillea,35(5):101-103,p1.86
261
De Toni, J.B. 1888. Sylloge Ustilaginearum et Uredinearum" in Sylloge
Fungorum, Saccardo P.A., Padova, Italy, 7:855
Ellis, M.B. 1976. More Dematiaceous Hyphomycetes. Commonw. Mycolog.
Inst., Kew, England
Howell, A. 1939. Studies on Histoplasma capsulatum and similar
form species. I. Morphology and development. Mycologia, 31:191-216
Mason, E.W. 1928. Annotated account of fungi received at the imperial
Boureau of Mycology. Myc. Pap., 2(1):37-39
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MYCOTAXON
VOIee RAV LL pps, 200-200 October-December 1986
ene PERSE
NOTES ON ENTOMOPHTHORALES (ZYGOMYCOTINA)
COLLECTED BY T.PETCH: II. ERYNIA ELLISIANA SP. NOV.,
NON ERYNIA FORFICULAE (GIARD), COMB. NOV.,
PATHOGENS OF FORFICULIDAE (DERMAPTERA)
ISRAEL S. BEN-ZE'EV
Department of Plant Pathology and Microbiology,
Faculty of Agriculture, The Hebrew University
of Jerusalem, P.0.Box 12, Rehovot 76100, Israel
ABSTRACT
Empusa forficulae var. major Petch 1944 (nomen nudum)
was reexamined and is redescribed here as an independent
species, Erynia ellisiana Ben-Ze'ev, sp. nov., characterized
by obovoid, uninucleate, bitunicate, subpapillate conidia
averaging 23.8 x 15.2 ym, simple or rarely sparsely digita-
te conidiophores, and lack of pseudocystidia and rhizoids.
Entomophthora forficulae Giard 1889 (=Zoophthora forficulae
(Giard) Batko 1964) is transferred to Erynia as E.forficulae
(Giard) Ben-Ze'ev, comb. nov. Both fungi are pathogens of
Forficulidae (Dermaptera) .
KEY WORDS: Entomophthorales; Erynia ellisiana; E.forficulae;
Dermaptera; Forficulidae; Forficula auricularia.
Petch (1944) described a fungus pathogenic on earwigs, Empusa for-
ficulae var. major Petch, as a variety of Entomophthora forficulae
Giard (1889), which he transferred to Empusa, as Empusa forficulae
(Giard) Petch. Since their original discovery, these two fungi were
seldom mentioned in the literature, and then only for classification
purposes, always quoting the original description (Guegen,1904 - quoted
in Petch, 1944; Lakon, 1919; Hutchison, 1963; Batko, 1964, 1966; Water-
house, 1975; Waterhouse and Brady, 1982). £E.forficulae Giard was appa-
rently reencountered only twice since its discovery: Rostrup (1893 -
quoted in Petch,1944) recorded it in Denmark, and a photograph showing
three European earwigs, Forficula auricularia L. killed by "Entomoph-
thora forficuli Giard" in the United States, is shown in Steinhaus
(1949) in his Fig. 102, without any further mention of this species in
the text. The description of £E.forficulae var. major in Waterhouse
(1975) and again in Waterhouse and Brady (1982) added new data unmen-
tioned by Petch (1944), on the existence and shapes of secondary coni-
dia and resting spores. These additions were based on a personal com-
munication from N. Wilding to G. Waterhouse (G. Waterhouse, pers. com-
mun. )
The fungus described by Petch (1944) was collected by E.A. Ellis in
England and resembled Giard's E.forficulae in having earwigs as hosts
264
and in lacking rhizoids. According to Petch, this fungus difered from
E.forficulae morphologically, in two ways: its conidiophores were simple
whereas those of E.forficulae were described as sparingly branched, and
its conidia were ca. 2.5 times wider, while their length was in the same
range as those of E.forficulae. According with the common practice of
classification at the level of species in the Entomophthorales, the two
fungi should have been considered as different species on morphological
grounds. However, Petch (1944) overemphasized the importance of the
host, as he did in several other instances, and as a result failed to
recognize new species, or mixed different species having a common host,
in one description (Ben-Ze'ev and Kenneth, 1981; Ben-Ze'ev, 1982, 1986).
As there is no specimen of Giard's £.forficulae left for examination,
one of Petch's specimens was reexamined in order to clarify the taxono-
mic position of E.forficulae var. major according to the modern classi-
fication of Entomophthorales, in which the old genus Entomophthora Fre-
senius had been split into five different genera, and some of its spe-
cies had been transferred to Conidiobolus Brefeld.
MATERIALS AND METHODS
The specimen examined, from the T.Petch collection, was kindly lent
by The Herbarium, Royal Botanical Gardens, Kew. It had no serial number
and its annotation was: "Empusa forficulae (Giard) Petch var. major
Petch n. comb., n. var., on earwigs, Old Lakenham, Norfolk. (Drawings
made on date of collection, E.A. Ellis)". These drawings were redrawn,
rearranged and provided with a scale, as Figs. 32-38 here.
Small pieces of fungal material were scraped with fine dissecting
needles from the outside and inside of the earwig's abdomen and were
further dissected, in drops of acetocarmine or acetoorcein, under a
dissecting microscope. The slides with the staining liquid and fungal
material were gently heated to remove air bubbles and were sealed with
coverslips. Measurements were taken with the aid of a calibrated eye-
piece, so that each division = 0.79 ym at a magnification of 40 x 25
(objective x ocular). The limit of resolution was less than 0.5 divi-
sion, < + 0.4 wm. Figures 1 - 31 were drawn with the aid of a camera
lucida.
RESULTS AND DISCUSSION
The specimen agreed, in macroscopic appearance, with the description
of Entomophthora forficulae by Giard (1889), with Fig. 102 in Steinhaus
(1949) and with Petch's (1944) description of Empusa forficulae var.
major. The conidiophores emerged and filled the spaces between adjacent
abdominal segments, and coalesced partially over some parts of the tho-
rax. Some conidial shapes (Figs. 1-26) agreed with the description in
Waterhouse (1975) and in Waterhouse and Brady (1982, no.65) and some
Figs. 1-38: Erynia ellisiana sp. nov.(=Empusa forficulae var. major
Petch): — 1-26, 32, 35-372, primany conidia: 30,43),, 358.54: hyphal
bodies; 27, 38: conidiophores, branched and unbranched, respectively;
28, 29: spherical bodies found in the type material — possibly imma-
ture resting spores, or structures unrelated to E.ellisiana; 1-31
drawn with the aid of a camera lucida at the scale shown at upper
right; 32-38 reproduced from original drawing by E.A. Ellis at the
scale shown at lower right.
265
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266
with Ellis' drawings (compare Figs. 2,5,6,22, and 24 with 32,35,36 and
37). Conidiophores were in very poor condition, but appeared to be
mostly unbranched as described by Petch (1944) and like Fig. 38 (re-
drawn from Ellis' drawings). One structure was found (Fig. 27) which
closely resembled sparingly-branched conidiophores of some Frynia spe-
cies, e.g. E.suturalis (Ben-Ze'ev, in preparation). It was difficult to
determine whether this structure was a three-branched conidiophore to
which another, simple one adhered, or a four-branched one. Mycelium
and hyphal bodies (Figs. 30,31) were also in very poor condition. Many
spherical, thin-walled bodies were present in the preparations (Figs.
28,29). These corresponded in size and yellow color with the resting
spores of E.forficulae var. major described in Waterhouse (1975) and in
Waterhouse and Brady (1982). However, several features seem to indicate
that these spherical bodies might not belong to the fungus under consi-
deration: a) the very thin walls and the numerous oily vacuoles filling
these cells would indicate young prespores, yet, in no instance were
these spherical bodies observed to be attached to hyphae or to hyphal
bodies, and no signs have been observed on their walls to indicate any
such attachment; b) the nuclei in the spherical bodies (Figs, 28,29)
were substantially smaller than those in conidia and hyphal bodies
(compare with other Figs.) Structures resembling mature entomophthora-
lean resting spores were not observed.
A comparison of conidial dimensions and other characters (Table 1)
shows that the fungus described as Entomophthora forficulae var. major
(Petch, 1944) is a species of Erynia Nowakowski emend. Humber § Ben-
Ze'ev (1981), and deserves an independent status as a new species:
ERYNIA ELLISIANA Ben-Ze'ev, sp. nov. (Entomophthoraceae)
=Empusa forficulae var. major Petch (nom. nud.) ,Trans. Br.
mycol. Soc. 27:87 (1944).
NON Entomophthora forficulae Giard, Bull. Sci. Fr. §& Belg.
20° 2114 GL880y,
CONIDIA PRIMARIA obovoidea, hyalina, uninucleata, bitunicata,
multivacuolata, plerumque subpapillata, raro epapillata aut
papillata {secundum Lakoni (1919) systema}, 19.8-30.0 x 12.6-20.2
um, med. 23.8 x 15.2 um. CONIDIA SECUNDARIA globosa, primariis
parviora, conidiophoris brevibus ex conidiis primariis portata.
CONIDIORUM NUCLEI entomophthoroidei, elliptici, 4.0-6.3 x 3.2-5,5
um, raro globosi, 4.0-6.3 um diametro. CONIDIOPHORA PRIMARIA
simplicia vel raro ramosa, determinata, ad 15 um lata. SPORAE
PERDURANTES, PSEUDOCYSTIDIA et RHIZOIDEA non observata.
In Forficulidae (Dermaptera) in Britannia.
TYPUS: Exsiccati et lamina per microscopium, Erynia ellisiana Ben-
ze'ev 1986 (=Empusa forficulae var. major Petch) designata, Reg.
Bot. Herb., Kew, Britannia.
PRIMARY CONIDIA obovoid, hyaline, uninucleate, bitunicate, multiva-
cuolate, mostly subpapillate, the rest epapillate or papillate f{acc. to
Lakon's (1919) classification}(Figs. 1-26 and 32,35-37) 19.8-30.0 x 12.6
~20 2m) (X= 23.08) x05) 2am, is= 72 2x1 OSepm n= SO), witha enero,
diameter ratio of 1.3-2.1 (X= 1.58, s= 0.17, n= 50). The papillar pro-
portional length (as % of total conidial length, epapillate conidia
excluded) is 3.1-18.9% (most frequently 6-8%); papillar proportional
width is 30-56.3% (m.f. 35-45%). SECONDARY CONIDIA were not observed in
this study, but have been reported (Waterhouse, 1975) to be smaller than
primary ones, globose, on short secondary conidiophores borne on primary
267
Table 1. A comparison of morphological characters of Entomophthorales
recorded as attacking Forficulidae (Dermaptera).
PR Tue ALR GY, CONIDIA
Length x Diam. Papillar
(min. - max.) Type
SPECIES X (um) CONIDIOPHORES
ent iis Renerhny (Dram yin een teem ty)
OSE Ratio Cytoplasm
Entomophthora 20-25 x 6-8 base hardly sparingly
forficulae 22x distinct branched
LER oh: a a a ae ara from apex ___
Forficula completely
auricularia L. Disc without
vacuoles
E.forficulae 18-21 x 8-10 5
(Rost rupli SOS ivestul9 59% 9 i
Sa erence ss adie po Bvt pontine Mahe felt SONNET 1) q
? ? ?
Empusa forficu-~ 20-30 x 15-18 subpapillate or
lae var. major ? papillate (epa-
Petch, 1944 Diriate acc, (to unbr anched
MET EAE fay Upc et lg oh BLES es al
earwigs ? multivacuolate
ACG aitoubllis*
drawings
Erynia 19.8-28.4 x 12.6-20.6 mostly
ellisiana Ben- PARTE aes Bon unbranched,
Ze'ev (=Empusa mostly rarely
forficulae var. subpapillate sparingly
major Petch) some papillate branched
CES PO Reap UEDA end TepApEEnaue
ete RR Peay ee | multivacuolate,
g Anas) uninucleate
i ee
conidia.
The lateral projections shown here on primary conidia (Figs,
17,18) could be such secondary conidiophores or ordinary germ-tubes.
CONIDIAL NUCLEI are entomophthoroid, with granular heterochomatin which
stained reddish-brown with acetoorcein and acetocarmine.
The stain
reaction was slower than in fresh material and apparently slower than
in other preserved material, probably due to poor preservation. Nuclear
shape was mostly ellipsoid, 4.0-6.3 x 3.2-5.5 wm, a smaller proportion
were globose, 4.0-6.3 um in diameter.
38) simple, rarely sparingly digitate, up to 15 um wide.
PRIMARY CONIDIOPHORES (Figs. 27,
PSEUDOCYSTIDIA,
RHIZOIDS and RESTING SPORES have not been observed in this study.
Pathogen of earwigs (Dermaptera: Forficulidae) in England.
TYPE: Specimen in T. Petch's collection annotated: Eryni ellisiana Ben-
Ze'ev 1986 (=Empusa forficulae var. major Petch) and microscope slides
annotated similarly, deposited at The Herbarium, Royal Botanical Gardens,
Kew, U.K.
268
This species is named in honour of Mr. E.A. Ellis, who collected it
along with many other British entomogenous fungi described or studied
by" Ts Petch.
When the genus Zoophthora Batko became a synonym of Erynia Nowakow-
ski, most species of the former were transferred to the latter (Humber
and Ben-Ze'ev, 1981; Ben-Ze'ev and Kenneth, 1982). Entomophthora forfi-
culae was overlooked — the following new combination is therefore
proposed:
ERYNIA FORFICULAE (Giard) Ben-Ze'ev, comb. nov.
basionym: Entomophthora forficulae Giard, Bull. Sci. Fr. § Belg.
20:211 (1889).
synonym: Zoophthora forficulae (Giard) Batko, Bull. Acad. Polon.
Sci. , 96r. oc)... Bi0l .cliidwi22404 Clo 6dj.
The fungus recorded by Rostrup (1893, cited in Petch, 1944) on ear-
wigs in Denmark (2nd in Table 1) appears to be a third, different spe-
cies attacking earwigs.
For the time being, and until £.forficulae and FE.ellisiana are
reencountered and reexamined from fresh material and more subgeneric
characters are described, it would be unsafe to decide to which subgenus
of Erynia they belong, although their unbranched or sparingly branched
conidiophores seem to indicate a possible affinity with the subgenus
Furia (Batko) Li §& Humber (1984).
ACKNOWLEDGMENTS
I thank the Curator of The Herbarium, Royal Botanical Gardens, Kew,
U.K. for the loan of E.forficulae var. major material, and Ms Grace
Waterhouse for valuable correspondence. I am grateful to Prof. Robert
G. Kenneth (The Hebrew University of Jerusalem) for his constructive
review of the manuscript, and to Prof. E.D. Kolman (Tel-Aviv University)
for the Latin description.
REFERENCES
BATKO, A.(1964). Some new combinations in the fungus family Entomophtho-
raceae (Phycomycetes). Bull.Acad.Polon.Sci.,Ser.Sci.Biol.cl II,
12:403-406.
BATKO, A.(1966). On the subgenera of the fungus genus Zoophthora Batko
1964 (Entomophthoraceae). Acta Mycol. 2:15-21.
BEN-ZE'EV, I.(1982). Erynia neopyralidarum sp. nov. and Conidiobolus
apiculatus, pathogens of pyralid moths, components of the misdescri-
bed species, Entomophthora pyralidarum (Zygomycetes: Entomophthora-
les). Mycotaxon 16:273-292.
BEN-ZE'EV. I.(1986). Notes on Entomophthorales (Zygomycotina) collected
by T. Petch: Erynia anglica comb. nov. and Erynia coleopterorum.
Mycotaxon 25:1-10.
BEN-ZE'EV, I. and KENNETH, R.G.(1981). Zoophthora radicans and Zoophtho-~
ra petchi sp. nov. (Zygomycetes: Entomophthorales), two species of
the 'sphaerosperma group" attacking leaf-hoppers and frog-hoppers
(Homoptera). Entomophaga 26:131-142.
BEN-ZE'EV, I. and KENNETH, R.G.(1982). Features-criteria of taxonomic
value in the Entomophthorales: II. A revision of the genus Erynia
Nowakowski 1881 (=Zoophthora Batko 1964). Mycotaxon 14:456-475.
GIARD, A.(1889). Sur quelques types remarquables de champignons entomo-
phytes. Bull.Sci.Fr. & Belg. 20:197~224.
269
HUMBER, R.A. and BEN-ZE'EV, I.(1981). Erynia (Zygomycetes: Entomophtho-
rales): emendation, synonymy, and transfers. Mycotaxon 13:506-516.
HUTCHISON, J.A.(1963). The genus Entomophthora in the Western Hemisphe-
re. fTrans.Kansas Acad.Sci. 66:237-254.
LAKON, G.(1919). Die Insektenfeinde aus der Familie der Entomophthoreen.
Z.Angew.Entomol. 5:161-216.
LI, Z. and HUMBER, R.A.(1984). Erynia pieris (Zygomycetes: Entomophtho-
raceae), a new pathogen of Pieris rapae (Lepidoptera: Pieridae) :
description, host range, and notes on Erynia virescens. Can.J.Bot.
62:653-663.
PETCH, T.(1944). Notes on entomogenous fungi. Trans.Br.mycol.Soc.
27 :81-93.
STEINHAUS, E.A.(1949) .'PRINCIPLES OF INSECT PATHOLOGY' Ist edition (ed.
E.A. Steinhaus) McGraw-Hill Book Comp.Inc.,New York, Toronto, London
WATERHOUSE, G.M.(1975). Key to the species Entomophthora Fres.
Bull.Br.mycol.Soc. 9:15-41.
WATERHOUSE, G.M. and BRADY, B.L.(1982). Key to the species of Entomoph-
thora sensu lato. Bull.Br.mycol.Soc. 16:113-143.
NOTE ADDED IN PROOF
Since this article was accepted, it was brought to my attention that
Erynta forficulae produces capilliconidia and belongs, therefore, to
Erynta subgen. Zoophthora (Balazy, S.,1984 — Sbornik Dokl. simpoz. po
teme KNTS-12U: Izucenie entomopatogennych microorganizmov, razrabota
technologii i primenenija. Rytro-Poznan:141-157; and pers. commun.).
The combination Erynta (Zoophthora) forficulae, used in that publication
is not, however, effectively or validly published.
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NOUmEA AVAL |b DD 0220 = 2.82 October-December 1986
CANOPARMELIA, PARAPARMELIA and RELICINOPSIS,
THREE NEW GENERA IN THE PARMELIACEAE (LICHENIZED ASCOMYCOTINA)
JOHN A. ELIX, JEN JOHNSTON and DOUGLAS VERDON
Chemistry Department, The Faculties, Australian Nattonal Untversity,
G.P.0. Box 4, Canberra, A.C.T. 2601, Australia
ABSTRACT: The genera Canoparmelia Elix and Hale gen.
nov., Paraparmelia Elix and Johnston gen. nov. and
Relictnopsts Elix and Verdon gen. nov. are here
segregated from Pseudoparmelia s. str. The main
differences between the genera lie in the chemical
properties of the upper cortex and the medulla, the
centres of distribution, the substrate and habitat
requirements and the size of the conidia and spores.
Introduction
The lichen genus Pseudoparmelia was erected by Lynge in 1914 on the
basis of a single collection from Brazil (Lynge 1914) and distinguished
from Parmelia sens. lat. by the presence of “pseudocyphellae” on the
lower surface. Santesson (1942) subsequently showed that the pseudo-
cyphellae were artefacts caused by the rhizines being torn off the lower
surfaces which revealed patches of the medulla.
More recently, Hale has accepted this typification of the genus
(Hale 1974, 1976) (Type species: Pseudoparmelta cyphellata Lynge) and
included in it all species previously accommodated in Parmelta subg.
Parmelta section Cyclocheila (Hale and Kurokawa 1964) or Parmelia sub-
gen. Cyclocheitla (Krog & Swinscow 1979). The genus so defined could be
recognized by the presence of simple rhizines and sublinear to irregul-
arly widened, often apically suhrotund, eciliate lobes. The apothecia
were adnate to substipitate with imperforate discs and the upper cortex
consisted of palisade plectenchyma with a pored epicortex.
Even so, it had become increasingly obvious that Pseudoparmelia so
defined, was a heterogeneous assemblage of species (Hale 1974, 1976).
As a result of further development and refinement of the generic con-
cepts within the Parmeliaceae (Hale 1986) it is apparent that Pseudo-
parmelta sens. str. only includes a small group of species closely
related to P. cyphellata Lynge: namely two other South American spec-
ies, P. hypomilta (Fée) Hale and P. chapadensis (Lynge) Hale, plus as
the more widely distributed lichen, P. sphaerospora (Nyl.) Hale. All
contain the same medullary pigment (secalonic acid A) while the upper
cortex contains only this pigment and atranorin (which is not accompan-
ied by chloroatranorin). Further, the spores are small and nearly
spherical (6 x 9 um), the conidia are elongated bifusiform to filiform
and the lower surface varies from a characteristic olivaceous brown to
pale brown in colour, with sparse concolorous rhizines. This combinat-
ion of characters clearly distinguishes Pseudoparmelta sens. str. from
the remaining species previously included in this genus (Hale 1974,
1975, 1986). Very recently Hale (1986) has separated the (Pseudo-
Zhe
parmelia) caperata group and transferred it to a new genus, Flavo-
parmelta.
During our revision of Pseudoparmelta sens. lat. in Australasia it
was soon apparent that there were four segregate groups previously
accommodated in Pseudoparmelta which are now excluded by the new circum-
scription of the genus. These were the P. tntertexta group (Reltcinop-
sits), the P. scotophylla group (Paraparmelia) and the P. texana groups
(Canoparmelta, containing most of the species accommodated previously in
this genus). These segregates differ from Pseudoparmelia sens. str. by
a combination of morphological, distributional, ecological, cortical and
chemical characters which have been summarized in Tables 1 and 2 and
discussed below.
Morphology
The spores and conidia
While the overall morphology of the apothecia and spores are
relatively uniform throughout the Parmeliaceae, the size of the spores
is quite variable. For example, Parmotrema and Flavoparmelta (the
Pseudoparmelia caperata group) consistently possess larger spores
(greater than 14 ym long) than do other Parmelta sens. lat. segregates
(less than 14 pm). By comparison Pseudoparmelia sens. str. has much
smaller (4-7 x 6-9 Um), almost spherical spores as does Reltctnopsts
(ellipsoid, 3-5 x 5-8 um). Analogous regularities in the size of spores
are observed for Canoparmelia with smaller spores than Flavoparmelia,
but larger than Paraparmelta, Pseudoparmelia and Reltctnopsts; and
Paraparmelia, which has spores smaller than Canoparmeltia, but larger
than those of Pseudoparmelta and Relictnopsts.
In several recent circumscriptions of parmelioid genera, consider-
able importance has been attributed to variations in the shape of the
conidia (Krog 1982a). For example, Punctelia is the only parmelioid
genus to consistently have unciform conidia while the related genus
Flavopunctelta has bifusiform conidia (Krog 1982b, Hale 1984).
The conidia of Pseudoparmelia sens. str. are markedly elongated (12-
20 um) by comparison with those of allied genera (Table 1) and vary from
elongated, weakly bifusiform to filiform in shape (Krog 1982a, b). At
the other extreme, Reltctnopsts and Paraparmelta are distinguished by
their consistently short (5-7 ym) cylindrical and bifusiform conidia
respectively. Flavoparmelta and Canoparmelia exhibit conidia of inter-
mediate length (7-10 um).
Thalline characters
Other major morphological distinctions between the segregates of
Pseudoparmelia sens. lat. as well as the consistency of characters among
the members of a particular segregate are apparent.
One such feature is the underside of the thallus. In Pseudoparmelta
sens. str. the lower cortex always is brown to olivaceous or pale brown
with concolorous rhizines which are always short, simple, and moderately
dense to the lobe margins. In Relicitnopsts the lower surface also is
pale tan to brown in colour and rhizinate to the margins but the rhiz-
ines are dense, elongated and entangled, and in R. intertexta densely
branched. The lower cortex of Canoparmelia, on the other hand, is black,
with a narrow, naked, brown marginal zone, and simple concolorous rhiz-
ines which vary from sparse to dense. In Paraparmelta the colour of the
lower surface varies from pale brown to jet black, and the rhizines are
short, moderate to sparse, stout and occasionally tufted at the apices.
The occurrence of vegetative propagules amongst the segregates of
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VITAWYVdOGNASd dO SALVOAYNDAS
T ATAVL
274
Pseudoparmelia also is noteworthy. All genera include isidiate species,
but sorediate and/or pustulate species are restricted to Flavoparmelia,
Canoparmelia, and Paraparmelia. Fully 50% of the species of Flavo-
parmelta and 40% of Canoparmelia exhibit such propagules, but they are
completely unknown in Pseudoparmelia sens. str. and Relictnopsis.
The lobe configuration is quite distinctive in two of the segregate
groups. In Relictnopsts the lobes are typically sublinear-elongate and
dichotomously branched, with the crowded or intricate lobe configuration
forming a thallus with overall gross morphology similar to that of many
species of Relitcitna. However, species of the latter genus are readily
distinguished by the presence of the highly characteristic bulbate cilia
and bifusiform conidia, whereas species of Reltcinopsis are eciliate and
have cylindrical to weakly fusiform conidia.
Although the lobe configuration is more variable in Canoparmelta,
most species have relatively broad (3-5 mm wide), irregular lobes with
rotund or subrotund apices. In Paraparmelta most species have relativ-
ely narrow (1-2 mm wide), sublinear to irregular lobes with subrotund
apices. Small foliose or subcrustose thalline growth forms also are
quite common in this genus.
Distribution
All three new genera have well defined centres of distribution.
Canoparmelta has its highest number of species in the Americas and
Africa, with some of the more widespread species, including C. texana,
C. carolintana, C. carneopruinata, C. crozalstana and C. salacinifera,
being essentially pantropical. Reltcinopsts, on the other hand, is re-
stricted to south-east Asia, the Indonesian archipeligo, Australia and
the Indian subcontinent with one report from Ivory Coast in tropical
Africa. Paraparmelia has two major centres of distribution both in the
Southern Hemisphere, (i.e. South Africa and Australasia) and is rare
outside these regions.
Ecology and Habitat
Although each of the genera exhibit some variability in their eco-
logical requirements, overall habitat differences are very obvious.
Paraparmelia is confined to saxicolous substrates in the dry
savannah woodland, subarid and arid areas of South Africa and Austral-
asia. In marked contrast, Relictnopsis is always corticolous and
restricted to the canopy and branches in tropical and montane rain-
forests. Most species of Canoparmelia also are corticolous, but some
species, such as C. caribaea are primarily saxicolous. Canoparmelta
species predominate in moist tropical and subtropical forests and wood-
land, and in this respect resemble Pseudoparmelia sens. str.
Chemistry
Cortical Chemistry
Despite the fact that Krog (1982 a,b) maintained that cortical
chemistry is generally unimportant above species level in the Parmel-
iaceae, we beg to differ. For instance, Esslinger (1977, 1978) found
that the distinctive cortical chemical reactions in the brown Parmeliae
were an excellent means for differentiating Melanelia, Allantoparmelia
and Neofuscelia. Indeed, Hale (1984) recently reinterpreted the generic
characters in the case of Punctelta (cortical atranorin) and Flavopunc-
telta (usnic acid) and concluded that the cortical chemical differences
in this case were mirrored by a number of significant morphological
(conidial) and biogeographical differences. It is important to re-
275
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276
iterate that no experimental or genetic proof has been introduced to
deny the potential value of cortical chemistry as a discriminator. As
Hale (1984) pointed out, a number of other lichen genera also are based
partly on highly correlated cortical chemistry e.g. Bulbothrix Hale
(atranorin) and Relicina (Hale & Kurokawa) Hale (usnic acid); Bryoria
Brodo & Hawksw. (brown pigments) and Alectoria Ach. (usnic acid).
Cortical chemistry is particularly useful for distinguishing the
segregates of Pseudoparmelia sens. lat. and is thus consistent with
Hale's belief that the nature and morphology of the cortex constitute
important taxonomic characters at the generic level (Hale 1981, 1984).
The upper cortex of all species of Paraparmelta contain atranorin
rarely in conjunction with chloroatranorin (by h.p.l.c.). Only in
P. capnoides, P. fwnigata, P. interposita and P. schistacea are small
amounts of usnic acid reported to occur (Kurokawa 1985) with atranorin,
giving these grey species a slight yellowish tinge on the apical lobes.
In Relictnopsis and Flavoparmelia the main cortical substance is usnic
acid (occasionally with small amounts of atranorin) giving these species
a distinct yellow or yellow-green colour.
The species of Canoparmelita invariably contain substantial quant-
ities of both atranorin and chloroatranorin in the cortex while Pseudo-
parmelia sens. str. contains small quantities of atranorin together with
the yellow medullary pigments, secalonic acid A and derivatives. The
latter pigments give the upper surface of Pseudoparmelta sens. str. a
dull yellowish cast. Several species of Canoparmelia (C. ecaperata, C.
zambiensts) consistently contain sufficient quantities of usnic acid
together with the major cortical substances, atranorin and chloroatran-
orin to colour the thallus yellow.
Medullary Chemistry
The main classes of medullary substances encountered in the species
of Pseudoparmelia sens. str., Canoparmelia, Paraparmelia and Relictnop-
Sts are detailed in Table 2. Although there is a significant overlap in
medullary chemistry of the segregates, definite trends can be observed.
In Paraparmelia the B-orcinol depsidones (salazinic acid, hypostict-
ic acid etc.) are predominant, whilst the orcinol depsidones (e.g. nor-
lobaridone) and orcinol depsides (e.g. lecanoric acid) are less common.
In Canoparmelta the orcinol depsides divaricatic acid and lecanoric
acid are particularly common, and the f-orcinol depsidones (stictic
acid, salazinic acid, norstictic acid) occur frequently. Broad variat-
ions in medullary chemistry from biosynthetically primitive depsides,
such as lecanoric acid, to more advanced depsidones has been reported
previously in the Parmeliaceae. Such well-established and distant
genera as Parmotrema and Xanthoparmelta exhibited parallel variations in
medullary chemistry so the range of metabolites recorded for Canopar-
melita and Paraparmelta are not inconsistent with these generic con-
cepts. The f-orcinol depsidones also are most common in both Pseudo-
parmelia sens. str. and Relicinopsts, but barbatic acid (a B-orcinol
depside) also is found in the latter. No orcinol depsides or depsidones
have been observed in these genera, although a number of unknowns
recorded for Pseudoparmelia sens. str. are not yet identified.
Paraparmelta_and Xanthoparmelia (Vainio) Hale
In Australia and South Africa, species of both Paraparmelia and
Xanthoparmelia (Vainio) Hale are well developed on rocks in subarid
areas, and exhibit striking morphological parallels. Hale (Hale 1976)
has interpreted these morphological similarities as representing an
277
example of convergent evolution of the genera in these stressed envir-
onments. Even though there are parallels in overall morphology, eco-
logical requirements and substrate preferences, we consider that the
differences between the taxa warrant their segregation at the generic
level.
Paraparmelta and Xanthoparmelia are clearly distinguished by cort-
ical chemistry (atranorin + chloroatranorin in Paraparmelta; usnic acid
in Xanthoparmelta) although a small minority of species of Paraparmelia
are known to develop pale yellow apical lobes due to the presence of
traces of usnic acid. Even so, atranorin is the preponderant cortical
metabolite in this genus and the upper surface of the lobes have a
typical mineral-grey colour. Usnic acid is preponderant in the charact-
eristic, yellow green upper cortex of the Xanthoparmeliae, but minor
traces of atranorin may be detected occasionally. The significance of
cortical chemistry has been discussed previously. The medullary-
chemical profiles of these two genera also show significant differences
- thus the B-orcinol depsides occur relatively frequently in Xantho-
parmelta (ca. 10% of species in Australasia) but are unknown in Parapar-
melita.
These two genera exhibit clear differences in world distribution.
While Paraparmelia is confined largely to the Southern Hemisphere (South
Africa, Australasia), Xanthoparmelia is widely distributed in both hemi-
spheres with centres of speciation in North America (ca. 25% species) as
well as Australia and South Africa. Paraparmelia and Xanthoparmelia
also show some significant variations in substrate preference and
thallus morphology. Although both genera are primarily saxicolous, many
Xanthoparmeltae (20% of the 117 species in Australia) are obligately
terricolous and commonly exhibit revolute or convolute lobes. Parapar-
melta has not been observed to be terricolous, and is not known to
exhibit a revolute- or convolute-lobed growth form.
Discussion
The main morphological, chemical, ecological and distributional
‘traits of the species in Pseudoparmelia sens. str., Relicinopsis and
Paraparmelia as well as the related segregates are outlined in Tables 1
and 2. They indicate that these segregates differ in a number of taxon-
omically important characters including spore size, conidial size and
morphology, habitat preferences, cortical chemistry, centres of distri-
bution, nature of vegetative propagules as well as several minor char-
acters. Since relative uniformity can be demonstrated within each of
the segregate groups, it provides substantial support for the proposed
generic splits.
Canoparmelia Elix & Hale gen. nov.
Thallus foliaceus, pagina superior cinerea, pagina inferior nigra,
rhizinas simplices instructa. Apothecia laminalia vel submarginali,
discis integris, sporis octonis, ellipsoideis, 10-14x6-8 um. Pycnidia
laminalia vel raro marginalia, conidiis bifusiformibus vel fusiformibus
vel cylindricis 7-10 um longis et cortex superior atranorinum et chloro-
atranorinum continens.
Typus generis: Canoparmelia texana (Taylor) Elix & Hale.
Thallus foliose, lobes subrotund at the apices, (0.5)-3.0-5.0
(-8.0) mm broad, eciliate. Upper surface uniformly grey, with a per-
forate polysaccharide covering. Medulla white. Underside black, rhiz-
inate, rhizines simple; with a narrow, paler, erhizinate zone at the
278
margins. Apothecia laminal or submarginal, disc entire, spores ellip-
soid, simple, 8 per ascus, 10-14x6-8 pm. Pycnidia laminal or rarely
marginal, conidia bifusiform more rarely cylindrical or weakly fusiform.
The cortex containing atranorin and chloroatranorin + usnic acid.
This new genus is considered to include the following species:
Canoparmelia adspersa (Vain.) Elix & Hale, comb. nov.
Basionym: Parmelita adspersa Vain., Hedwigia, 46, 168. 1907.
Canoparmelia amazonica (Nyl.) Elix & Hale, comb. nov.
Basionym: Parmelia amazonica Nyl., Flora, Jena 68, 611. 1885.
Canoparmelia aptata (Krempelh.) Elix & Hale, comb. nov.
Basionym: Parmelita aptata Krempelh., in Nylander, Fora, Jena 52,
291. \:1869%
Canoparmelia caribaea (Hale) Elix & Hale,, comb. nov.
Basionym: Parmelia caribaea Hale in Hale & Kurokawa, Contrib. U.S.
Nat. Herb., 36, 152. 1964.
Canoparmelia carneopruinata (Zahlbr.) Elix & Hale,, comb. nov.
Basionym: Parmelia carneopruinata Zahlbr., Sitzungs. kaiser. Akad.
Wissen., Math.-Natur. Klasse, 1, 419. 1902.
Canoparmelia caroliniana (Nyl.) Elix & Hale, comb. nov.
Basionym: Parmelia caroliniana Nyl., Flora, Jena 68, 614. 1885.
Canoparmelia cinerascens (Lynge) Flix & Hale, comh. nov.
Basionym: Parmelta ctnerascens Lynge, Arkiv Bot., 30B, 104. 1914.
Canoparmelia concrescens (Vain.) Elix & Hale, comb. nov.
Basionym: Parmelta conecrescens Vain. in F. Welwitsch, Cat. African
Plants Collected by F. Welwitsch in 1853-61, 2, 400. 1901.
Canoparmelia corrugativa (Kurok. & Filson) Elix & Hale, comh. nov.
Basionym: Parmelia corrugattva Kurok. & Filson, Bull. Natn. Sci.
Mus.,), lokyo,0B els 138.5) 1975.
Canoparmelia crozalsiana (Bouly de Lesd.) Elix & Hale, comb. nov.
Basionym: Parmelia crozalstana Bouly de Lesd. ex J. Harmand, Lichens
de France, 4, 555. 1910.
Canoparmelia cryptochlorophaea (Hale) Elix & Hale, comb. nov.
Basionym: Parmelta cryptochlorophaea Hale, Bryologist, 62, 18. 1959.
Canoparmelia ecaperata (Mull. Arg.) Elix & Hale, comh. nov.
Basionym: Parmelia ecaperata Mull. Arg., Flora, Jena 74, 378. 1891.
Canoparmelia epileuca (Hale) Elix & Hale, comb. nov.
Basionym: Parmelia epileuca Hale, Bryologist, 75, 343. 1972.
Canoparmelia eruptens (Kurok.) Elix & Hale, comb. nov.
Basionym: Parmelia eruptens Kurok., in Hale & Kurokawa, Contrib.
U.S. Nat. Herb., 36, 153. 1964.
Canoparmelia inhaminensis (Dodge) Elix & Hale, comb. nov.
Basionym: Parmelia inhaminensis Dodge, Ann. Missouri Bot. Gard., 46,
130.37 1959
Canoparmelia inornata (Hale) Elix & Hale, comb. nov.
Basionym: Parmelta inornata Hale, Phytologia, 22, 32. 1971.
Canoparmelia martinicana (Nyl.) Elix & Hale, comb. nov.
Basionym: Parmelta martintcana Nyl., Flora, Jena 68, 609. 1885.
Canoparmelia nairobiensis (Stein. & Zahlbr.) Elix & Hale, comb. nov.
Basionym: Parmelia natrobiensis ("“neirobiensis”) Stein. & Zahlbr. in
Zahlbr., Bot. Jahrb. Syst., 60, 517. 1926.
Canoparmelia norsticticata (Stevens) Elix & Hale, comb. nov.
Basionym: Parmelta norsticticata Stevens in Elix & Stevens, Aust. J.
Bote L277 8815091979.
Canoparmelia raunkiaeri (Vain.) Elix & Hale, comb. nov.
Basionym: Parmelta raunkiaert Vain., Ann. Acad. Scien. Fennicae, 6A,
19 LOTS
279
Canoparmelia salacinifera (Hale) Elix & Hale, comb. nov.
Basionym: Parmelta salacinifera Hale in Hale and Kurokawa, Contrib.
UsSs Nat. Herb. , 36, °157: 1964.
Canoparmelia schelpei (Hale) Elix & Hale, comb. nov.
Basionym: Parmelta schelpet Hale, Bryologist, 75, 344. 1972.
Canoparmelia scrobicularis (Krempelh.) Elix & Hale, comb. nov.
Basionym: Parmelia scrobicularis Krempelh., Vid. Medd. Natur. For.
Kj6benhaven, 25, 10. 1873.
Canoparmelia somaliensis (Mull. Arg.) Elix & Hale, comb. nov.
Basionym: Parmelia somaliensts Mull. Arg., Flora, Jena 68, 501.
1885.
Canoparmelia subtiliacea (Nyl.) Elix & Hale, comb. nov.
Basionym: Parmelia subtiliacea Nyl., Flora, Jena 68, 614. 1885.
Canoparmelia texana (Tuck. ) Elix & Hale, comb. nov.
Basionym: Parmelia texana Tuck., Amer. J. Sci. Arts, ser. 2, 25,
424, 1858.
Canoparmelia zambiensis (Hale) Elix & Hale, comb. nov.
Basionym: Parmelta zambiensis Hale, Bryologist, 75, 346. 1972.
Canoparmelia zimbabwensis (Hale) Flix & Hale, comb. nov.
Basionym: Parmelia zimbabwensis Hale, Bryologist, 75, 346. 1972.
Paraparmelia Elix & Johnston, gen. nov.
Thallus foliaceus, saxicola, adnatus vel laxe adnatus, lobis sub-
irregularibus, marginibus eciliatis, paginae superiore epicorticato
cinerea, atranorinum continens, subtus pallide brunneus vel niger,
modice vel sparse rhizinosus, rhizinis simplicibus. Apothecia adnata,
imperforata, sporis octonis, ellipsoideis, 4-6 x 7-10 um. Pycnidia
laminalia, conidiis bifusiformibus, brevibus, 5-7 wm longis. Typus
generis: Paraparmelta scotophylla (Kurok.) Elix & Johnston.
Thallus foliose, saxicolous, adnate to loosely adnate, lobes sub-
irregular, narrow (0.5) 1.0-3.0 (-4.0) mm wide, margins eciliate. Upper
surface grey, often darkening with age, with a perforate polysaccharide
covering. Medulla white. Underside pale tan to jet black, moderately
to sparsely rhizinate, rhizines simple for the most part, sometimes
tufted at the apices. Apothecia adnate, imperforate, spores ellipsoid,
simple, 8 per ascus, 7-10 x 4-6 ym. Pycnidia laminal or subapical,
conidia bifusiform. The cortex containing atranorin, rarely chloro-
atranorin or traces of usnic acid.
This genus currently contains the following species:
Paraparmelia adusta (Kurok.) Elix & Johnston, comb. nov.
Basionym: Parmelta adusta Kurok., Bull. Natn. Sci. Mus., Tokyo, B,
MAR Ta L985 6
Paraparmelia alabamensis (Hale & McCull.) Elix & Johnston, comb. nov.
Basionym: Parmelta alabamensts Hale & McCull., Bryologist, 71, 44.
1968.
Paraparmelia annexa (Kurok.) Elix & Johnston, comb. nov.
Basionym: Parmelia annexa Kurok., in Hale & Kurokawa, Contrib. U.S.
Nat. Herb., 36, 151. 1964.
Paraparmelia arcana (Kurok.) Elix & Johnston, comb. nov.
Basionym: Parmelita arcana Kurok. in Hale and Kurokawa, Contrib. U.S.
Nat. Herb., 36, 151. 1964.
Paraparmelia astricta (Brusse) Elix & Johnston, comb. nov.
Basionym: Parmelia astricta Brusse, Bothalia, 15, 315. 1984.
Paraparmelia basutoensis (Hale) Elix & Johnston, comb. nov.
Basionym: Parmelia basutoensis Hale, Bryologist, 75, 342. 1972.
280
Paraparmelia capnoides (Kurok.) Elix & Johnston, comb. nov.
Basionym: Parmelia capnotdes Kurok., Bull. Natn. Sci. Mus., Tokyo,
Beh) Veer LooD.
Paraparmelia cerussata (Kurok.) Elix & Johnston, comb. nov.
Basionym: Parmelta cerussata Kurok., Bull. Natn. Sci. Mus., Tokyo,
BATS hai9est
Paraparmelia condyloides (Kurok.) Elix & Johnston, comb. nov.
Basionym: Parmelia condyloides Kurok. in Hale, Bryologist, 75,
S43. 19 12%
Paraparmelia conranensis (Elix) Elix & Johnston, comb. nov.
Basionym: Parmelta conranensts Elix, Aust. J. Bot., 29, 20. 1981.
Paraparmelia fumigata (Kurok.) Elix & Johnston, comb. nov.
Basionym: Parmelta fumigata Kurok., Bull. Natn. Sci. Mus., Tokyo, B,
11 8101985"
Paraparmelia interposita (Kurok.) Elix & Johnston, comb. nov.
Basionym: Parmelia interpostta Kurok., Bull. Natn. Sci. Mus., Tokyo,
Byallt81. wtoRSs
Paraparmelia ischnoides (Kurok.) Elix & Johnston, comb. nov.
Basionym: Parmelia tschnoides Kurok. in Hale and Kurokawa, Contr.
U.S. Nat. Herb., 36, 155. 1964.
Paraparmelia lecanoracea (Mull. Arg.) Elix & Johnston, comb. nov.
Basionym: Parmelia lecanoracea Mull. Arg., Flora, Jena 71, 529.
1888.
Paraparmelia lithophila (Kurok.) Elix & Johnston, comh. nov.
Basionym: Parmelta lithophtla Kurok., Bull. Natn. Sci. Mus., Tokyo,
Boa lise 63.471985..
Paraparmelia lithophiloides (Kurok.) Elix & Johnston, comb. nov.
Basionym: Parmelia lithophtloides Kurok., Bull. Natn. Sci. Mus.,
Tokyo, By, 1ig.e4s. 1985.
Paraparmelia molybdiza (Nyl.) Elix & Johnston, comb. nov.
Basionym: Parmelia molybdiza Nyl., in Crombie, J. Bot. Brit. For.,
TARAS 76:
Paraparmelia mongaensis (Elix) Elix & Johnston, comb. nov.
Basionym: Parmelta mongaensis Elix, Aust. J. Bote, 29, 22. 1981.
Paraparmelia mrina (Kurok.) Elix & Johnston, comb. nov.
Basionym: Parmelia murina Kurok., Bull. Natn. Sci. Mus., Tokyo, B,
Ligkes wages:
Paraparmelia nana (Kurok.) Elix & Johnston, comb. nov.
Basionym: Parmelta nana Kurok., Bull. Natn. Sci. Mus., Tokyo, B, l,
86. 1985.
Paraparmelia neoquintaria (Hale) Elix & Johnston, comh. nov.
Basionym: Parmelta neoquintaria Hale, Smithonian Contr. Rot., 31,
S598 970%
Paraparmelia numinbahensis (Elix) Elix & Johnston, comb. nov.
Basionym: Parmelta numinbahensis Elix in Elix & Stevens, Aust. J.
BOts;% 27 Ook 97 9.6
Paraparmelia owariensis (Asahina) Elix & Johnston, comb. nov.
Basionym: Parmelta owartensis Asahina, J. Jap. Bot., 28, 135. 1953.
Paraparmelia prolata (Hale) Elix & Johnston, comb. nov.
Basionym: Parmelia prolata Hale, Bryologist, 75, 344. 1972.
Paraparmelia pustulescens (Kurok.) Elix & Johnston, comb. nov.
Basionym: Parmelta pustulescens Kurok. in Hale and Kurokawa, Contr.
U.S. Nat. Herb., 36, 156. 1964.
Paraparmelia rodriguesiana (Hue) Elix & Johnston, comb. nov.
Basionym: Parmelia rodriguestana Hue, Nouv. Arch. Mus. Paris, ser.
SeeuGMilore: 99.
281
Paraparmelia rupicola (Lynge) Elix & Johnston, comb. nov.
Basionym: Parmelia ruptcola Lynge, Akiv Bot., 13, 132. 1914.
Paraparmelia schistacea(Kurok. & Filson) Elix & Johnston, comb. nov.
Basionym: Parmelta schistacea Kurok. & Filson, Bull. Natn. Sci.
Mus., Tokyo, B, 1, 44. 1975.
Paraparmelia scotophylla (Kurok.) Elix & Johnston, sp. nov.
Basionym: Parmelia scotophylla Kurok. in Kurokawa and Filson, Bull.
Natn. Sci. Mus.', Tokyo, B, 1, 45. 1975.
Paraparmelia spodochroa (Kurok. & Filson) Elix & Johnston, comb. nov.
Basionym: Parmelita spodochroa Kurok. & Filson, Bull. Natn. Sci.
MUS.) WLOKVO. Bo. 1s 4005 ol 9/5,
Paraparmelia subtortula (Hale) Elix & Johnston, comb. nov.
Basionym: Parmelia subtortula Hale, Phytologia, 27, 3. 1973.
Paraparmelia tortula (Kurok.) Elix & Johnston, comb. nov.
Basionym: Parmelia tortula Kurok. in Hale and Kurokawa, Contr. U.S.
Nat. Herb., 36, 157. 1964.
Paraparmelia vanderbylii (Zahlbr.) Elix & Johnston, comb. nov.
Basionym: Parmelia vanderbylii Zahlbr., Ann. Cryptog. Exot., 5
2520 L932
Paraparmelia violacea (Kurok.) Elix & Johnston, comb. nov.
Basionym: Parmelta violacea Kurok. in Hale and Kurokawa, Contr. U.S.
Nat. Herb., 36, 158. 1964.
Paraparmelia xanthomelaena (MUll. Arg.) Elix & Johnston, comb. nov.
Basionym: Parmelta xanthomelaena Mull. Arg., Flora, Jena, 66, 48.
1883.
Relicinopsis Elix & Verdon, gen. nov.
Thallus foliaceus, corticola adnatus, lobis sublinearibus vel
linearibus, margines eciliatis, cortice superiore epicorticato,
viridolutescens, acidum usnicum continens, subtus pallide brunneus vel
brunneus, dense rhizinosus, rhizinis elongatis, simplicibus vel
furcatis, concoloris. Apothecia adnata, imperforata, sporis octonis,
ellipsoideis, 5-8 x 3-5 pm. Pycnidia marginalia raro laminalia, plus
minusue extrusus, conidiis cylindricis vel raro fusiformibus, brevibus
5-7 um longis.
Typus generis: Relictnopsts intertexta (Mont. & v.d. Bosch) Elix &
Verdon.
Thallus foliose, corticolous, lobes sublinear to linear-elongate,
narrow, 1.0-2.0 (-3.0) mm wide, with eciliate margins. Upper surface
yellow-green, with a perforate polysaccharide covering. Medulla
white. Underside pale tan to brown, densely rhizinate to the margins,
rhizines elongate, simple or irregularly branched, concolorous with the
lower surface. Apothecia adnate, imperforate, spores ellipsoid, simple,
8 per ascus, 5-8 x 3-5 pm. Pycnidia marginal rarely laminal, sometimes
extruded, conidia cylindrical or fusiform. The cortex containing usnic
acid and accessory atranorin.
This new genus includes four species:
Relicinopsis dahlii (Hale) Elix & Verdon, comb. nov.
Basionym: Pseudoparmelia dahlit Hale, Smithsonian Contrib. Bot., 31,
28. 1976.
Relicinopsis intertexta (Mont. & v.d. Bosch) Elix & Verdon, comb. nov.
Basionym: Parmelta intertexta Mont. & ved. Bosch in Montagne,
Sylloge Gen. Spec. Cryptog., 327. 1856.
282
Relicinopsis malaccensis (Nyl.) Elix & Verdon, comb. nov.
Basionym: Parmelia malaccensis Nyl. in Nylander and Crombie, J.
Linn. Soc. London, 20, 52. 1883.
Relicinopsis rahengensis (Vain.) Elix & Verdon, comb. nov.
Basionym: Parmelia rahengensis Vain., Ann. Soc. Zoo0l.-Bot. Fenn.
Vanamo nl oo. Loz.
Acknowledgements
We wish to thank the Australian National University for generous
financial assistance through the Faculties Research Fund and Dr. M.E.
Hale for his continual encouragement, helpful and stimulating discuss-
ions, and assistance with type material.
References
Esslinger, T.L. (1977) A chemosystematic revision of the brown
Parmeliae. Journ. Hattori Bot. Lab., 42, 1-211.
Esslinger, T.L. (1978) A new status for the brown Parmeltae. Mycotaxon,
7, 45-54.
Hale, ME. Jr. (1974). New Combinations in the Lichen Genus
Pseudoparmelia Lynge. Phytologia, 29, 188-191.
Hale, M-E. Jr. (1976). A monograph of the Lichen Genus Pseudoparmelia
Lynge (Parmeliaceae). Smithsonian Contrib. Bot., 31, 1-62.
Hale, M.E. Jr. (1981). Pseudocyphellae and pored epicortex in the
Parmeliaceae: their delimitation and evolutionary significance.
Lichenologist 1, 1-10.
Hale, M.E. Jr., (1986). Flavoparmelia, a new Genus in the Lichen Family
Parmeliaceae (Ascomycotina) Mycotaxon, 25, 603-605.
Hale, M.E. Jr.e, and Kurokawa, S. (1964). Studies on Parmeltia subgenus
Parmelia. Contrib. U.S. Nat. Herb., 36, 121-191.
Krog, H. (1982a). Evolutionary trends in the foliose and fruticose
lichens of the Parmeliaceae. Journ. Hattori Bot. Lab., 52, 303-311.
Krog, H. (1982b). Punctelta, a new lichen genus in the Parmeliaceae.
Nord. J. Bots.) 2,267 —292
Krog, H. and Swinscow, T.D.V. (1979). Parmelia subgenus Hypotrachyna in
East’ Africas’ Norw. J.’ Bot. ,/26, (11-43.
Kurokawa, S. (1985). Studies on Australian and Tasmanian Species of
Parmelta (2), Bull. Natn. Sc. Mus., Tokyo, B, 11, 77-90.
Lynge, B. (1914). Die Flechten der ersten Regnellschen Expedition. Die
Gattungen Pseudoparmelia gen. nov. und Parmelia Ach. Arkiv for
Botanik, 13) 2-172.
Santesson, R. (1942). Pseudoparmelia Lynge, a Lichen Genus to be
rejected. Svensk Botanisk Tidskrift, 36, 471-474.
Vobis, G. and Hawkswork, D.L. (1981). Conidial Lichen-Forming Fungi in
"Biology of Conidial Fungi" Vol 1, Ch 9, pp 245-272 (Academic Press,
New York).
MYCOTAXON
Vole XXVIT, “pp. 283-288 October-December 1986
TWO NEW DISCOMYCETES ON PINUS
A. FUNK
Pacific Forestry Centre
Canadian Forestry Service
506 W. Burnside Road
Victoria, B.C.
V8Z 1M5
ABSTRACT
Two new inoperculate discomycetes are described from
Pinus in North America, viz., Claussenomyces pini sp.
nov. from P. contorta Dougl., and Crumenulopsis
lacrimiformia sp. nov. from P. palustris Mill.
INTRODUCTION
Collections of discomycetes received from widely
separated locations in North America have presented an
opportunity to describe two new bark-inhabiting species
on Pinus. The first is from north-central British
Columbia, Canada on lodgepole pine (Pinus contorta
Dougl.) and the second is from central Florida, U.S.A. on
longleaf pine (Pinus palustris Mill.). Both fungi are
associated with disease in pine stems but their
pathological status is not known.
TAXONOMY
Claussenomyces pini sp. nov. Fig. 1-4
Anamorphosum: Dendrostilbella-similis in agaro cultae
Apothecia atra, nitida, sessilia vel substipitata,
Singularia vel subcaespitosa, marginata, circularia vel
undulata, ionomidotica, 0.2-0.8 mm diametro, usque ad 0.4
284
mm alto. Excipulum ex hyphis radiata composita; hyphis
atro-viridia, gelatinosa, irregularia, 2-4 um lata.
Asci clavati, longistipitati, octospori, apices
incrassatis, poro in iodo non caerulescenti, 85-105 x
8-10 um. Ascosporae hyalinae, fusoideo-ellipsoideae,
rectae vel leniter curvatae, 1-3 septatae, 10-18 x 3
um, raro in ascis ascoconidia productae, sed in agaro
ascoconidia numerosae. Ascoconidia hyalina, continua,
ovoidea, 4 x 2 um. Paraphyses filiformes, ramosae, ad
apices leniter incrassata, epithecium fuscum formantibus.
Holotypus: in caulibus Pinus contorta, Houston, B.C.,
Canada, January 1985, DAVFP 23301.
Apothecia black, shiny, sessile to substipitate, single
or sub- caespitose, marginate, circular or undulate,
ionomidotic, 0.2-0.8 mm diameter, up to 0.4 mm high.
Excipulum composed of radiating hyphae; hyphae dark
green, gelatinized, irregular, 2-4 wm diameter. Asci
clavate, long-stalked, 8-spored, thickened apex, pore not
staining in iodine, 85-105 x 8-10 wm. Ascospores
hyaline, fusoid-ellipsoid, straight or slightly curved,
1-3 septate, 10-18 x 3 um, rarely producing ascoconidia
in the ascus, but abundantly on agar. Ascoconidia
hyaline, continuous, ovoid, 4 x 2 um. Paraphyses
filiform, branching, slightly swollen at the tips and
forming a dark epithecium.
Holotype: in a stem of Pinus contorta Dougl. collected 20
km west of Houston, B.C., Canada in January 1985 by W.G.
Bristow. DAVFP 23301.
Claussenomyces pini apothecia were found at the edge of a
stalactiform rust canker (Cronartium coleosporioides
Arth.) in January and were in prime condition. It would
appear to be a secondary invader of diseased bark in this
Single known occurrence. The asci when mounted in dilute
Figs. 1-4: Claussenomyces pini 1. Apothecia, habit on
pine bark. X 88. 2. Asci and ascospores. XK 572. 3.
Asci mounted in dil. KOH showing extension of inner
contents. K 572. 4. Germinating ascospores with
ascoconidia. XK 572. Figs. 5-8: Crumenulopsis
lacrimiformia 5. Apothecia, habit on pine bark. KX 88.
6. Asci and ascospores. X 572. 7. Apothecium, median
section. X 242. 8. Excipular tissues. KX 704.
285
5
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i
AN
l,
i,
) '
286
KOH extrude their contents in a manner resembling the
action normally found in bitunicate asci (Fig. 3).
Cultural study
Cultures were readily obtained from ascospores shot from
rehydrated apothecia onto malt agar. Ascospores
germinated first by production of large quantities of
ascoconidia giving the young colony a moist, yeast-like
appearance. Ascospores and ascoconidia (Fig. 4)
eventually produced germ tubes and the mycelial colony
developed. Colonies were entirely appressed, sodden,
almost colorless and with no diffusion zone. Optimum
growth temperature was approximately 20°C at which the
colony expanded 2.5 cm in 3 weeks. A strong antibiosis
was noted against common mold contaminants in the
plates. In 5 weeks the colony became yellowish brown and
small synnemata developed. The synnemata were at first
concolorous but later darkened with a hyaline droplet of
conidia at the tip; they measured approximately 0.5 mm
high and 0.5 mm wide. Conidia were rod-like, continuous
and 2 x 0.8 wm.
Crumenulopsis lacrimiformia sp. nov. (Figs. 5-8)
Apothecia erumpentia, singularia vel caespitosa, atro
brunnea, circularia vel undulata, substipitata,
gelatinosa, rugosa, hymenium flavida, 0.6-1.1 mm
diametro. Excipulum medullosum subhyalina, textura
epidermoidea; excipulum ectalum atro brunnea, usque ad
20um crassum, textura angularis. Asci clavati,
octospori vel quadrispori, poro in iodo non
caerulescenti, 65-85 x 10-11 wm. Ascosporae hyaline,
non septatae vel raro uniseptatae, guttulatae,
lacrimiformae, 13-17 x 4-5 um. Paraphyses simplices,
filiformes, epithecium non formantibus, 2 wm crassa, ad
apicem leniter incrassata.
Holotypus: DAVFP 23300, in rami Pinus palustris Mill.
Gainesville, Florida, U.S.A., Aug. 11, 1985, J.D. Rogers.
Apothecia erumpent, single or caepitose, dark brown,
circular or undulated, substipitate, roughened,
gelatinized, hymenium pale yellow, 0.6-1.1 mm diameter.
Medullary excipulum subhyaline, composed of textura
epidermoidea; ectal excipulum dark brown, approximately
287
20 um thick, composed of textura angularis. Asci
clavate, eight-spored or four-spored, pore not staining
in iodine, 65-85 x 10-11 um. Ascospores hyaline,
non-septate or rarely one-septate, tear-drop shaped,
13-17 x 4-5 um. Paraphyses simple, filiform, not
forming an epithecium, approximately 2 um wide,
slightly swollen at the tips.
Holotype: DAVFP 23300, on dead tips of Pinus palustris
Mill. near Gainesville, Florida, U.S.A., 11 Aug. 1985,
collected by J.D. Rogers.
Isotype: WSP 67857.
DISCUSSION
The genus Claussenomyces Kirschst. was characterized as
having green to black, gelatinized apothecia with J-asci
and septate ascospores that "bud" in the ascus (Korf
1973; Dennis 1978). Dennis (1956) gave full descriptions
of some species under the name Corynella Boud. which Korf
and Abawi (1971) showed to be invalid. The new species
appears to be closest to C. prasinulus (Karst.) Korf and
Abawi (1971), having ascospores of similar size and
septation, but C. prasinulus has emerald green apothecia
and paraphyses that branch repeatedly at the tip.
Another species on pine, C. dacrymycetoideus Ouellette
and Korf (1979), has 7-21l-septate ascospores that bud
prolifically in the ascus. It is possible that
ascospores of the new species might bud more profusely in
the ascus if allowed to mature further. Because the
collection was made in mid-winter it is conceiveable that
budding would recommence with the return of higher
temperatures.
The genus Claussenomyces is also represented in British
Columbia by C. pseudotsugae (Groves) Ouellette and
Pirozynski (1974), a species occurring on Pseudotsuga,
and by C. olivaceus (Fckl.) Sherwood (Hawksworth and
Sherwood 1981) a resinicolous species on conifers. These
two species have pycnidial anamorphs and are otherwise
easily distinguished from C. pini on apothecial
characters.
Crumenulopsis lacrimiformia was found on dead tips on
pine branches in central Florida. No cultures were
288
made. It is closest to C. pinicola (Fries) Groves
(Groves 1969) but it is smaller throughout and has
distinctive tear-drop shaped ascospores. The genus
Crumenulopsis was placed in the family Dermateaceae by
Korf (1973) and it was he who first suggested that this
fungus might be a new species of Crumenulopsis. The
structure and habit are very close to the other two
species in the genus (Groves 1969) which also occur on
pine.
ACKNOWLEDGEMENTS
I am indebted to Drs. J.D. Rogers and R.P. Korf for the
Crumenulopsis collection and to Wen-ying Zhuang for
preparing microsections; Dr. J.A. Parmelee kindly loaned
collections from Herb DAOM; Mr. Leo Unger surveyed pine
plantations for Clausssenomyces.
Dennis, R.W.G. 1956. A revision of the British
Helotiaceae in the herbarium of the Royal Botanic
Gardens, Kew, with notes on related European
species. Commonweal. Mycol. Inst. Pap. No. 62, Kew.
Dennis, R.W.G. 1978. British ascomycetes. J. Cramer,
Vaduz.
Groves, J.W. 1969. Crumenulopsis, a new name to replace
Crumenula Rehm. Can. J. Bot. 47: 47-51.
Hawksworth, D.L. and M.A. Sherwood. 1981. A reassess-
ment of three widespread resinicolous discomycetes.
Can. J. Bot. 59: 357-372.
Korf, R.P. 1973. Discomycetes and Tuberales. in The
Fungi, an advanced treatise 4A: 249-319. Academic
Press, New York.
Korf, R.P. and G.S. Abawi. 1971. On Holwaya, Crinula,
Claussenomyces, and Corynella. Can. J. Bot. 49:
1879-1883.
Ouellette, G.B. and K.A. Pirozynski. 1974. Reassessment
of Tympanis based on types of ascospore germination
within asci. Can. J. Bot. 52: 1889-1911.
Quellette, G.B. and R.P. Korf. 1979. Three new species
of Claussenomyces from Macaronesia. Mycotaxon 10:
255-264.
MYCOTAXON
VoIRaxkVCLy opp atZ89=319 October-December 1986
A Comparison Between Phytophthora cryptogea and P. drechsleri
H. H. Ho and S. C. Jong
Department of Biology, State University of New York, New Paltz,
N.Y. 12561; Mycology & Botany Department, American Type
Culture Collection, 12301 Parklawn Drive, Rockville, MD 20852
Abstract
Considerable controversies exist in literature concerning the
identification of a plant pathogen as Phytophthora cryptogea or P.
drechsleri. Phytophthora drechsleri is believed to be distinguished.
from P. cryptogea primarily by its ability to grow well at 35 C and
secondarily, by its occasionally homothallic behavior, larger
oogonia and larger, more elongate sporangia with tapered base.
Present study was undertaken to compare 19 isolates of P.
cryptogea with 11 isolates of P. drechsleri (including the type
sadick authentic cultures) from various hosts all over the world.
Based on morphological and physiological characters, most isolates
could not be recognized as "typical" P. cryptogea or P.
drechsleri; they possess certain diagnostic features of both
species. Growth at 35 C, alone or in combination with other
criteria, could not separate these isolates into two distinct
species. It appears most practical at this point, to merge P.
drechsleri with P. cryptogea which has priority.
Introduction
Phytophthora cryptogea Pethybr. & Laff. is a common plant
pathogen in the temperate regions attacking over 80 plant species
from 23 families (45). First described on tomato (50), it is
characterized by small hyphal swellings in clusters, ovoid to
obpyriform, non-papillate, internally-proliferating sporangia,
amphigynous antheridia and relatively small spherical oogonia.
Tucker (68) recognized that the Phytophthora isolate from rotting
potato tuber was similar to P. cryptogea but differed in the ability
to grow well at 35 C, and thus a new species, P. drechsleri was
erected. Later, Waterhouse (71) added that P. drechsleri could be
further differentiated from P. cryptogea by the occasionally
homothallic behavior, narrow hyphae, larger oogonia and larger, more
elongated sporangia with tapered base. However, in practice,
researchers often encountered great difficulty in deciding whether
an isolate should be named P. cryptogea or P. drechsleri because the
temperature criterion might not correlate with the morphological
characters or the isolate has characteristics intermediate between
these two species (18, 22, 27, 34, 41). Consequently, there was
considerable confusion in literature concerning the identity of a
pathogen being P. cryptogea or P. drechsleri. For example, the
290
pathogen on sunflower, cucurbits and potato tubers, respectively,
has been identified by some as P. cryptogea (35, 56, 57) but as P.
drechsleri by the others (4, 14, 68). A researcher often reported
both species associated with the same disease, as on Transvaal daisy
(66), safflower (34), Douglas-fir (53) walnut (40) and cherry (43,
73). Nearly half of the plant species attacked by P. drechsleri
are also hosts for P. cryptogea (9, 45, 64). Bumbieris (6
demonstrated that isolates of P. cryptogea and P. drechsleri had
similar growth rate, hyphal diameter and sporangial shape and size
and suggested the merging of these two species under P. cryptogea.
The proposal has received support (12, 23, 26, 27, 41, 62) while
some prefer to keep them as separate species (37, 40, 43, 53, 73).
In view of the uncertainty, Keim et al. (33) identified the pathogen
on Washington palm as belonging to P. drechsleri/cryptogea complex.
Present study was undertaken to compare isolates of P. cryptogea
and P. drechsleri from a variety of hosts all over the world to
determine if these two species should be treated as synonymous or
separate taxa.
Materials and Methods
Isolates and media. Specific information on the isolates of
P. cryptogea and P. drechsleri used is given in Table 1. For
crossing experiments, P. cinnamomi 32992 (A2), 32993 (Al), P.
almivora 26200 (A2), 26201 (Al) and P. nicotianae 38606 (A2) and
38607 (Al) were used. All isolates were obtained from the American
Type Culture Collection (ATCC), Rockville, Maryland. Culture media
used in various experiments included cleared V-8 agar (CV8), Difco
cornmeal agar (CMA), and fresh corn meal (CA) agar as used by
Tucker (68).
Table 1. Isolates of Phytophthora cryptogea and P. drechsleri used
Species ATCC Host Origin Source
P. cryptogea 15402 Aster sp. Unknown Gallegly N57
=CBS 308.62
15403 Godetis sp. S. Africa Gallegly N56
=CBS 307.62
28193 Pinus radiata S. Australia Bumbieris P10
soil
28194 Pinus radiata S. Australia Bumbieris P30
soil
34301 Pseudotsuga Oregon Pratt 37
menziesii
36226 Unknown S. Australia Halsall $121
96229 Eucalyptus Tasmania, Halsall D128
forest Australia
36301 Potato Ohio Rowe 116
44672 Garden Soil S. Australia Gerrettson-
Corne!l $121
46518 Chrysanthemum California MacDonald
morifolium Ph13
46721(A) Aster sp. USA Zentmyer
P1088
=(BS 270.39
Table l.
Species
P. cryptogea
P. drechsleri
(Continued)
ATCC
46999
48234
48637
52401
52402
52403
52404
56962(T)
10923
15404
15428
26756
28200
34302
44388
44389
46723
46724(A)
56192
58424
58425
58597
Host
Grapevine
Juniperus
chinensis
Unknown
Begonia-
Elatior
hybrid
Solanum
marginatum
Begonia-
EBlatior
hybrid
Begonia
Elatior
hybrid
Lycopersicon
esculentum
Unknown
Citrus
Dalmatian
insect flower
Carthamus
tinctorius
Safflower
Douglas fir
Pigeon pea
Pigeon pea
Pinus radiata
Beta vulgaris
var. altissima
Cucumis sativa
Cucumis
Lycopersicon
esculentum
Unknown
sativa
Origin
S. Africa
California
Unknown
W. Germany
Ecuador
W. Germany
W. Germany
Ireland
USA
California
Unknown
Unknown
Australia
Oregon
India
India
California
USA
China
Iran
Japan
Japan
291
Source
Marais 4
MacDonald 2-B
Warner
Kroeber
63651
Kroeber
63689
Kroeber
63787
Kroeber
64132
HotAiz.t
=CBS°113-.19
Jeffers
Gallegly N112
=CMI 40532
Gallegly Nlll
=CMI 40499
Gother
CMI 164.185
Stovold GES 1
Roth 47
Erwin P2
Erwin P4
Zentmyer
P1076
Zentmyer
P1087
=(5B5,292.59
FOS nee
Flos inten
= Ershad
rOtital bes oe
= Miyata
LER Slt
Ho, (tio 36
a A0 ed |
ATCC-Identified by American Type Culture Collection accession number
CBS -Centraalbureau voor Schimmelcultures, Baarn, Netherlands
CMI -Commonwealth Mycological Institute, Kew, Surrey, England
A -Authentic culture
T -Type culture
292
Morphology: Colony characteristics on CV8, CMA, and CA were
compared ‘after incubating in darkness at 20 C for 7 days. The
colony diameters on CV8 were measured at right angle through the
inoculum and the width of primary hyphae measured under light
microscope. The minimal and maximal temperatures for growth were
tested by growing them on CV8 and CMA at 5 C and 35 C. Sporangia
were produced by incubating small mycelial agar discs of CV8 in
freshly collected stream water sterilized by filtration through 0.45 um
pore size millipore membrane discs and incubating under light at
20 C. Production of sex organs in single cultures was followed on
CV8 supplemented with sitosterol (30 mg/l) by examining them
periodically under microscope through the bottom of the petri dish.
If the isolate failed to produce sex organs readily in single
culture, it was paired with the appropriate mating types of P.
cinnamomi, P. cryptogea, P. drechsleri, P. nicotianae or a
palmivora. In case of a successful mating, the ability of the
isolate to produce sex organs by selfing was confirmed by pairing it
with the compatible strain across a polycarbonate membrane to
prevent physical contact between the cultures (36). They were
examined for sex organs after 3-4 wk incubation in dark at 20 C. To
induce chlamydospore formation, small mycelial agar discs were
submerged in 15 ml sterile distilled water in 19 x 150 mm test
tubes, and incubated in dark at 15 C for 5 wk.
Ability to grow on malt extract agar: Isolates were grown on
Difco malt extract solidified with 1.5% Bacto agar. Colony
diameters were measured after 6 days.
Pigment production: Isolates were grown on Timmer's medium
(55). Production of pigment in the medium was determined after 2 wk
at 20 C in the dark.
Growth response to malachite green: Malachite green was
added to CMA at a concentration of 1:18,000,000. Colony diameters
were measured after 6 days.
Ability to utilize nitrate nitrogen: Isolates were grown on
Ribeiro's synthetic agar medium (55) minus asparagine so that
nitrate nitrogen was the only nitrogen source. Colony diameter was
determined after 6 days.
Pathogenicity to apple: Apple fruit (McIntosh) was
inoculated through artifical wound which was then sealed with Scotch
tape and enclosed in a plastic bag. Results were determined after
incubation at 20 C for 2 wk.
Results
Colony morphology: Colony morphology of the isolates on CV8
ranged from slightly fluffy to fairly fluffy with no distinct growth
pattern. The edge of colony was slightly diffuse and irregular.
Colonies were mainly appressed on CMA. Most isolates of P.
cryptogea and about half of isolates of P. drechsleri produced
luxuriant aerial mycelium on fresh corn meal agar medium. The
primary hyphae on CV8 were uniform, measuring 5-7um wide, though
the branches might be uneven. Network or chains of small spherical
to irregular hyphal swellings (<25 um) were often produced in water,
or on membranes in crossing experiments.
BOS
Sporangium: Sporangia were produced readily in water within
24-48 hr by all isolates except P. cryptogea isolates ATCC 48234 and
ATCC 56962 which did not sporulate until 4-5 days later. In all
cases, the sporangia were non-deciduous and non-papillate and
internal prolifetion was seen at least in some sporangia. The apex
of sporangium often flattened on mounting and the empty sporangia
collapsed partially after zoospore release. Sporangia were mainly
obpyriform, ovoid to elongate, with tapered or rounded base in
varying proportions (Table 2). Repeated emergence of zoospores was
observed in ATCC 52403 and ATCC 52404 of P. cryptogea and ATCC
58597 of P. drechsleri. a
Sex organs: Despite numerous attempts, P. drechsleri isolate
ATCC 28200 did not produce any sex organs whereas ATCC 34302 only
mated with Al mating type of P. nicotianae to form nicotianae -type
oongia. The production of sex organs by other isolates are
summarized in Table 3. Oogonia were spherical to subspherical with
no tapered base, though the oogonial stalk was sometimes tapered
within the antheridium. Oospores were plerotic in all isolates
filling the oogonia to varying extent, except in the selfing of ATCC
15428 of P. drechsleri in old cultures. The free space between the
00goum wall and the oospore inside is represented by the following
formula:
Free oogonial space = Oogonium diam - oospore diam X 100 (%)
Oogonium diam
The thickness of oospore wall (2-5 um) relative to the
oospore size is expressed as follows:
Relative oospore thickness = Outer oospore diam - inner oospore
diam X 100
Outer oospore diam
In general, oogonia produced by selfing of heterothallic
isolates were larger, developed pigmentation earlier and more
intensely than those formed in crossings, but they often aborted,
devoid of properly developed oospores. In crossing with P.
cinnamomi, both large cinnamomi -type and smaller cryptogea/
drechsleri - type oogonia were produced. Antheridia were short
cylindrical, mostly unicellular. Bicellular antheridia were
occasionally found especially among isolates of P. drechsleri or in
crosses with P. cinnamomi.
Chlamydospores: No chlamydospores could be found in any
isolate of P. cryptogea or P. drechsleri.
Physiology: All isolates produced very slight pigmentation
in Timmer's medium except ATCC 15403 and ATCC 48637 of P.
cryptogea and ATCC 15428 of P. cryptogea which formed moderate
brown coloration. They grew well on malt agar medium, tolerated
malachite green and utilized inorganic nitrate readily. With the
exeption of ATCC 15404 of P. drechsleri, all isolates caused
lesions in apple fruit. a
294
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Discussion
For the sake of comparison, the overall means of sporangia
and sex organs of all P. cryptogea and P. drechsleri isolates used
in present study are summarized in Table 4 and the published data on
these two species in Table 5.
Tucker (68) erected P. drechsleri primarily based on its high
maximal temperature for growth. A review of the published
literature revealed that some isolates of P. cryptogea with
morphological characteristics typical for the species grew at or
above 35 C (6, 18, 34, 41, 54), and not all isolates of P.
drechsleri prefer high temperature for growth (6, 46, 53, 59). In
present study, out of 16 isolates of P. drechsleri five did not grow
at all and 3 showed only traces of growth at 35 C. On the other
hand, at least one isolate of P. cryptogea grew well and 3 isolates
had slight growth at this temperature. Phytophthora drechsleri is
supposed to stop growing at 10 C (68) or 5C 70) while the minimal
temperature for growth for P. cryptogea is under 1 C (71) or under 5
C (68). In present study, about half of the P. cryptogea isolates
failed to grow whereas 5 out of 14 isolates of P. drechsleri showed
good or marginal growth at 5 C. Such discrepancy also existed in
literature (5, 6, 34, 43, 53, 66). It is not clear if the isolates
used in this study have undergone any change since initial isolation
but if they did, it merely shows that the temperature threshold
difference may not be a stable taxonomic character.
Newhook et al. (44) distinguished P. cryptogea with broad
hyphae (>8um diam) from P. drechsleri with narrow hyphae (<8im
diam). Waterhouse (71) described the hyphae of P. cryptogea as very
uneven, up to 8ym and those of P. drechsleri as fairly uniform, not
often over 5 um wide. Present study demonstrated that the primary
hyphae of both species were mostly uniform, 5-7um wide. Similarly,
Bumbieris (6) found no significant difference in the hyphal width
between these two species. Published literature shows that the
hyphal width of P. cryptogea was under 8um, similar to P.
drechsleri. In fact, Waterhouse and Blackwell (72) described the
mycelium of P. cryptogea in the British Isles as fine (3 \ta 5 fm
wide).
One of the major reasons why Waterhouse (71) kept P.
cryptogea separate from P. drechsleri is that the former produces
smaller sporangia (37-40 ym long) compared with the larger sporangia
(36-50 um long) in the latter. We found that the overall sporangial
length of all P. cryptogea isolates studied is slightly but not
significantly smaller than that of P. drechsleri isolates (49 + 8 vs
52 + 10 um) because of the great variations in measurements,
especially among isolates of P. drechsleri. It is hard to compare
our results with the published data, because of the different
conditions under which they were produced, but the mean values of
the sporangial length were 30-62 um for P. cryptogea and 29-64im
for P. drechsleri ,(Table 5) suggesting again that the difference in
sporangial size is not significant enough to separate the two
species.
Most researchers consider the sporangia of P. drechsleri more
elongated than those of P.cryptogea (16). To certain extent, this
difference is confirmed in present study which shows the overall L/B
307
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ratio to be 1.7 + 0.2 for P. cryptogea isolates and 1.9 + 0.3 for P.
drechsleri isolates but the difference can hardly be adequate to
distinguish these two species, especially in view of the overlapping
ranges: 1.4-2.0 and 1.4-2.3 respectively. Both oval/obpyriform
type and elongated type sporangia were present in isolates of these
two species, but the latter type was more common in P. drechsleri
isolates. The overlapping ranges were also reflected in literature
by the average L/B ratios reported: 1.2-1.9 for P. cryptogea and
1.2-2.1 for P. drechsleri (Table 5). Although most of the sporangia
of P. drechsleri in Tucker's diagram (68) were elongated, this
feature was not mentioned in his species description and it is not
clear if elongated sporangia indeed characterized this species.
Based on Tucker's original data (68), the L/B ratio of sporangium in
P. drechsleri was 1.5, the same as that of P. cryptogea as given by
Pethybridge and Lafferty (50) and even less than that determined by
Tucker himself (1.7). Thus, it is unlikely that the elongated
sporangium was used by Tucker as a diagnostic character to
differentiate P. drechsleri from P. cryptogea. Waterhouse (71)
described the sporangia of P. drechsleri as "Very variable in shape
from broadly obpyriform to elongated obpyriform".
Wilcox and Mircetich (73) and Chitzanidis and Koyeas (8)
considered the tapering base of the sporangium as a key character in
distinguishing P. drechsleri from P. cryptogea which has rounded
base. Present study showed that while tapered base was more common
among isolates of P. drechsleri, it was also present in P.
cryptogea isolates s and many isolates produced sporangia with
rounded or tapered base in varying proportions. Both rounded and
tapered bases were figured by Tucker for P. drechsleri (68) while
Waterhouse (71) referrred to the sporangia of P. drechsleri as
"sometimes tapering at the base".
Waterhouse (71) noted that the sporangia of P. cryptogea
often has a conspicuous central vacuole. We have Cuescued such
vacuoles in many isolates of P. drechsleri and P. cryptogea and they
do not seem to be characteristic of P. cryptogea only. In fact, a
central vacuole is shown in every sporangium of P. drechsleri
figured by Tucker (68). Gerrettson-Cornell (23, 24) found central
vacuoles only in sporangia that failed to germinate by zoospores.
Another key character used by Waterhouse (71) and Krober (37)
to separate ie cryptogea from P. drechsleri is the oogonial size
which is smaller in the former (30-32 vs 36 um diam). The oogonial
size was not discussed when Tucker (68) erected P. drechsleri as a
new species. His measurement of oogonia of P. drechsleri (av. 31.3
um diam) was slightly larger but very close to that given by
Pethybridge and Lafferty (50) for P. cryptogea (av. 30m diam).
Other researchers reported the average oogonial diameters to be
20-40 um for P. cryptogea and 25-43um for P. drechsleri (Table 5)
In present study, the overall mean oogonial size of P. drechsleri
(33 + 3 um diam) was slightly but not significantly Targer than that
of ee cryptogea (31 + 3um diam) and the range of variation was the
same (28.5 -38um diam) for both species. Furthermore, when oogonia
formed by selfing alone or by crossing alone were considered there
was hardly any difference in the oogonial diameters between P.
cryptogea and P. drechsleri (34 + 3 vs 35 + 4 and 31 + 2 vs SH ONS) + 3
um diam, respectively). In general, as reported earlier in Re
cinnamomi (29), the oogonia produced by selfing in present study
were larger than those formed in dual cultures and they developed
pigmentation more rapidly and more intensely. It is important for
taxonomic studies to induce the formation of sex organs in single
isolate culture. Unfortunately, we found out that although a
heterothallic isolate mated readily with a compatible partner on
agar medium, it often failed to produce any or only a few fully
developed oogonia with oospores when they were separated physically
by a polycarbonate membrane. This was especially true for isolates
of P. cryptogea. It appears that while sex organs may be formed by
selfing when two compatible strains are crossed in dual cultures,
the possibility of hybridization cannot be dismissed (22, 63).
Newhook et al. (44) considered the oospore wall thickness of
. cryptogea relative to the oospore diameter a particularly
roan diagnostic character. Present study compared the
percentage of the oospore diameter occupied by the oospore wall and
found that the difference too small (43+5 % and 41+5 %,
respectively) and the range overlapping too much (29-51% and 27-47.5
% respectively) to differientiate P. cryptogea from P. drechsleri.
According to Waterhouse (71), the antheridia of P. cryptogea
are shorter than those of P. drechsleri (10 and 14-15 um,
respectively), but she and Blackwell (72) earlier described the
antheridia of P. cryptogea from the British Isles as "spherical, 16
m in diameter". No measurement of the antheridia was given in the
original descriptions of P. cryptogea (50) and P. drechsleri (68),
although Tucker described the antheridia of the latter as
"variable". Other reseachers reported the antheridia to be 12-19 um
long in P. cryptogea and 10-24m long in P. drechsleri. We found
the overall antheridia of P. drechsleri to be slightly longer than
that-otsri. cryptogea (15 + 1 and 14 + 1m, respectively) but again
the difference is too small and the range overlapped (13-16.5 and 13-
15.5 um, respecively) so that this character cannot be used as a
dependable taxonomic criterion. Although long, becellular
antheridia were more often found in isolates of P. drechsleri, the
occurrence was too erratic to be of significance in differentiating
these two species.
Waterhouse (71) further noted that whereas P. cryptogea is
heterothallic, producing oospores rapidly and abundantly with P.
cinnamomi, P. drechsleri may be homothallic or heterothallic ‘but
would not mate with P. cinnamomi. Our observation confirmed earlier
reports (31, 52, 58, 61, 62) that isolates of P. drechsleri could
mate with P. cinnamomi whereas some isolates of P. cryptogea failed
to produce sex organs a crossed with P. cinnamomi. It is
difficult to distinguish P. cryptogea and P. drechsleri based on
sexual behavior. Both species behaved heterothallically (58) and
although homothallic eae are more common for P. drechsleri they
have been Lebel ies for P. cryptogea as well (57). Leonian (3
distinguished P. cryptogea from P. drechsleri because the former
but not the latter produced sex organs from sterile mycelium when
transferred from pea broth to distilled water, implying homothallism
in P. cryptogea. In present study, all isolates were heterothallic
except ATCC 10923 of P. drechsleri. Phytophthora drechsleri
isolate ATCC 58424 produced sex organs in single culture within two
weeks but formed them more abundantly and more rapidly when mated
with the compatible strain. It is not known whether any of the
314
heterothallic isolates studied was homothallic when freshly
isolated, but some isolates of P. drechsleri from cucumber were
homothallic initially and then changed to heterothallic condition
(Ho, unpublished).
Although chlamydospores of P. drechsleri have been reported
on plant roots and hypocytyls (2,10,11), these structures could
not be found in culture, despite diligent search among all isolates
of P. cryptogea and P. drechsleri. Pal et al. (48) reported
chlamydospores produced in situ by P. drechsleri var. cajani, but
based on their diagrams, the structures considered as chlamydospores
should be interpreted as hyphal swellings because they were not cut
off from the rest of the mycelium.
In cultural characteristics, isolates of P. cryptogea and P.
drechsleri were very similar especially on CV8 and CMA.
Physiologically, they were indistinguishable in the ability to grow
on malt extract agar, to utilize inorganic nitrate, to tolerate
malachite green and in the pigment production. Pathologically, they
all caused lesions on apple fruits within two weeks. The only
exception, ATCC 15404 of P. drechsleri which failed to grow on apple
fruit despite numerous attempts, could have degenerated as it also
showed unusually slow growth compared with other isolates.
In summary, there is no reliable taxonomic character to
differentiate P. cryptogea from P. drechsleri. When the isolates
were plotted based on the most important criteria used: mean
sporangial length, mean oogonial diameter and growth at 35 C, it was
not possible to separate them into distinct cryptogea group nor
drechsleri group. The majority of isolates had certain
characteristics of both species. Thus, isolate ATCC 10923 of P.
drechsleri produced rather large oogonia abundantly and promptly in
single culture but formed smaller, obpyriform to ovoid sporangia
with rounded base and failed to grow at 35 C. Isolate ATCC 58424 of
P. drechsleri produced large, elongated sporangia with tapered
base, grew well at 35 C but the oogonia were small. Isolate ATCC
48637 of P. cryptogea produced small, obpyriform sporangia with
rounded base but grew well at 35 C and formed rather large oogonia.
It can be argued that some of these could be hybrids. On the other
hand, in view of the large number of intermediate forms, the most
practical solution at this point is to merge P. drechsleri with P.
cryptogea which has priority. The merging of these two species was
proposed more than 30 years ago by Bywater and Hickman (7) who
noticed the similarity between P. _pruptoges, P. drechsleri, P.
erythroseptica, P. himalayensis . erythroseptica sensu
Waterhouse) and P P. richardiae a suggested grouping them together
as one species: P. erythroseptica. Bumbieris (6) raised this issue
again by proposing to treat P. cryptogea and P. drechsleri as
conspecific. The similarity between the two species was further
substantiated by electrophoresis pattern of soluble protein (41) and
serology (26). Our study substantiated the claim that P.
drechsleri cannot be specificially differentiated from P.
cryptogea. We feel that one of the major problems __
in Phytophthora taxonomy is that there are too many species to be
distinguished by too few taxonomic characters which are too easily
affected by cultural conditions. At least, one way to solve the
$15
problem is to compare many isolates of all Phytophthora species
under standard conditions to determine if some of these "species"
could be consolidated. Although we do not subscribe to Leonian's
notion (38) that there are only three good species
of Phytophthora, Ribeiro's estimate (55) of "only a dozen or so
valid species" does not appear to be such a "radical view" either.
Differences between "species" can often be attributable to
adaptation, physiological specialization or hybridization, and
should be treated as morphological variation within a taxon. Thus,
although morphological differentiation existed in host-specialized
groups of P. megasperma (28), it was viewed only as a consequence
of isolation due to host specialization. A similar situation
probably exists here for P. cryptogea. Thus "P. drechsleri" can be
considered as a high temperature variant of P. cryptogea adapted to
plant hosts of warmer areas and along with this change, minor
modifications in morphological characteristics as well. We prefer
to refrain from a formal re-description of P. cryptogea until many
isolates of other related species are studied to determine if any
further regrouping might be necessary.
Acknowledgement
This work was supported in part by a grant from the SUNY New
Paltz Grants for Research and Creative Program to H.H. Ho and
NSFevorant BSI (9415523 sto. S.C. Jong.
The authors thank Li Luo and Laura Palmisans for their
invaluable technical assistance and Elmer E. Davis for reviewing the
manuscript.
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479-482. |
50. Pethybridge, G.H., and Lafferty, H.A. 1919. A disease of
tomato and other plants caused by a new species of
Phytophthora’. . Sci. Proc. RR.» DublinsSoceu(N7S.. 715s, 487-5055
51. Pratt, B.H., and Heather, W.A. 1972. Method for rapid
differentiation of Phytophthora cinnamomi from other
Phytophthora species isolated from soil by lupin baiting.
Trans Br... Mycol. *S0e< #593 °87-96.
52. Pratt, B.H., and Heather, W.A. 1973. Recovery of potentially
pathogenic Phytophthora and Pythium spp. from native vegetation
in Australia. Aust. J. Biol. Sci. 26: 575-582.
53. Pratt, R.G., Roth, L.F., Hansen, E.M., and Ostrofsky, W.D.
1976. Identity and pathogenicity of species of Phytophthora
causing root rot of Douglas-fir in the Pacific Northwest.
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54. Rattink, H. 1981. Characteristics and pathogenicity of six
Phytophthora isolates from pot plants. Neth. J. Pl. Pathol.
87: 85-90.
55. Ribeiro, O.K. 1978. A Sourcebook of the Genus Phytophthora.
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56. Rowe, R.C., and Schmitthener, A.F. 1977. Potato pink rot caused
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57. Satour, M.M., and El-Shinnawy, S.A. 1972. A new root and
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MYCOTAXON
VOM Sou Dn DDiieo 2 biti S eo October-December 1986
2 RE EA GN ATL NUE cdl 3 AE Sh a eT Sl a a a
TREMELLOSTEREUM (TREMELLACEAE) NOV. GEN.
by
L. RYVARDEN
Dep. tof, Biology, Div. of Botany, Univ...of Oslo, ‘P.0. Box
1045 Blindern, N-O316 Oslo 3, Norway.
SUMMARY
Tremellostereum nov. gen. is described with T. dichroum
(Lloyd) Ryv. as type species. It is characterized by a
stereoid fruitbody, covered by a cottony tomentum and
has cruciate basidia with 2 sterigmata.
In 1922 Lloyd described Stereum dichroum based ona
collection by Brace from The Bahamas. He suggested it
could be a degenerate form of Gloeoporus dichrous (Fr. )
Bres. although he stated that pores were completely
absent. He observed neither spores nor basidia. During an
examination of Lloyds species described as Stereum, I
discovered that S. dichroum has cruciate basidia with two
large sterigmata. Thus, it clearly belongs in the
Tremellaceae. A search in the literature (Lowy 1971 and
McNabb 1973) revealed that there was no suitable genus to
accomodate a species with such set of characters.
Tremellostereum (Tremellaceae) nova. gen.
Fructificatio annua, pileata, pileus albus ad ochraceus,
gossypinus, hymenium laevis, gelatinosus, roseus, systema
hypharum monomiticum, hyphae generatoriae afibulatae,
basidia longitudinalia septata, ovoidea cum 2
sterigmatibus grande, sporae ellipsoideae, hyalinae,
tenuitunicatae, non-amyloideae.
Type species: Tremellostereum dichroum (Lloyd) Ryv.
Fruitbody annual, pileate, sessile and dimidiate, soft
when fresh, light of weight and brittle when dry, pileus
white to ochraceous, cottony, spongy to fibrillose,
hymenium smooth, dense and cartilaginous when dry,
probably gelatinous and pink to pale red when fresh, pale
reddish brown when dry, hyphal system monomitic,
generative hyphae without clamps, basidia longitudinally
septate with two large sterigmata, spores ellipsoid,
smooth, hyaline, thinwalled and non-amyloid.
BVA
10 pm
m
Fig. 1. Tremellostereum dichroum A) section through
fruitbody, B) fruitbody seen from above, C) basidia, D)
spores, E) hyphae from the tomentum. From the holotype.
WAS
Tremellostereum dichroum (Lloyd) Ryv. comb. nov.
Basionym: Stereum dichroum Lloyd, Lloyd mycol. Writ.
Praltooes 1922.
Fruitbody annual, pileate, up to 5 cm long and 3 cm wide,
sessile and slightly dimidiate, probably flexible when
fresh, light of weight and brittle when dry, pileus white
to sand-coloured, cottony to fibrillose and spongy,
easily compressed, tomentum up to 4 mm deep, azonate to
slightly zonate, margin sharp, hymenium smooth, probably
pink to pale red and gelatinous when fresh, pale brown
and cartilaginous when dry, strongly contrasting the
light-coloured and cottony context.
Hyphal system monomitic, generative hyphae with simple
septa in the hymenium sinuous, interwoven, richly
branched, hyaline and thinwalled, 2-6 wm wide, in the
context and tomentum parallel, slightly thickwalled,
straight and with scattered septa.
Metabasidia ovoid, scattered in the hymenium, 20-30 x
8-10 um, hyaline or partly filled with oily inclusions,
longitudinally septate and with two large sterigmata,
12-32 x 3-6 pm, cystidia not seen, spores ellipsoid to
subcylindrical with one straight side, hyaline,
thinwalled and non-amyloid, 10-12 x 6-7 um,
spore-repetetion not observed.
Only the type is known: Bahamas, Leg. L.J.K. Brace, Lloyd
herb. no 8070 (BPI), on dead wood, causing a white rot
judged from the attached wood.
The new genus is characterized by its stereoid habit,
with a light, spongy and white context and
microscopically by its two-sterigmate basidia.
Microscopically the closest relative seems to he
Tremellodendron, which, however, has branched, erect
Clavaria-like fruitbodies and basidia with 4 sterigmata.
Microscopically the new genus and Tremellodendron share
the same type of simple septate hyphae and ellipsoid,
hyaline spores of approximately the same size.
Acknowledgements
Din bo LOWY, bousitana State University, has critically
read the manuscript and suggested improvements for which
I am very grateful.
References
Lioyd, C.G: 1922: Mycological Notes no.67. Lloyd Mycol.
Writ. 7:1137-1168. - Lowy, B. 1971: Treméllales, Flora
Neotropica Monograph no 6. New York. - McNabb, R.F.R.
1973: Phragmobasidiomycetidae: Tremellales,
Auriculariales, Septobasidiales, p. 303-316 in Ainsworth,
G.C., Sparrow, F.K. & Sussman, A.S. (eds.) The Fungi.
Vol. IVB. London.
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MYCOTAXON
VOIP ARVLES | DDs 452 51545 October-December 1986
Sn ENE ADRES bbe olin D2 RE RS ES Le Rill ahold ak a alan hs ase
BASIDIOMYCETES THAT DECAY GAMBEL OAK
IN SOUTHWESTERN COLORADO: III
J. Page Lindsey
Department of Biology
Fort Lewis College, Durango, Colorado 81301
SUMMARY
Sixteen additional Basidiomycetes that decay Quercus
gambelii Nutt. in southwestern Colorado are illustrated and
described. Two species are described as new: Hyphoderma
anasaziense and Platygloea laplata. A comprehensive key
includes all species described in this series of papers.
In two previous papers (Lindsey, 1985; Lindsey 1986) fifty
wood-rotting Basidiomycetes were described as decay agents of Quercus
gambelii Nutt., Gambel oak, in southwestern Colorado. This paper
continues and concludes this series with descriptions and illustrations
of an additional sixteen Basidiomycetes. Microscopic techniques are
described in the initial paper of the series (Lindsey, 1985). Numer-
ical color names are from Munsell’s Book of Color (Munsell, 1976);
descriptive color names are the author’s.
Annotated list of Additional Species that Decay Gambel Oak
ARMILLARTIA MELLEA (Vahl:Fr.) Quél., Champ. Jura et Vosges, p. 75.
1872.
Basidiocarps annual, centrally stipitate, single to caespitose around
stumps and living trees; pileus up to 15 cm diam, honey colored to
golden brown (7.5YR 3/6 to 7.5YR 5/6), covered with fine pointed
scales, these often washing off after rains; pileus margin inrolled at
first, expanding to plane, slightly striate; lamellae adnate to
decurrent, cream colored to light brown; stipe up to 15 cm long and 2.5
em diam, often with a _ slightly bulbous base, light brown; annulus
superior, whitish. Hyphal system monomitic; subhymenial hyphae
thin-walled, nodose-septate, the clamps often distorted or difficult to
see, 2.5-4 um diam; tramal hyphae thin-walled, nodose-septate but the
clamps often so skewed as to appear simple-septate, up to 8 um diam.
Cystidia none. Sterile, hyphoid, simple-septate hymenial elements
present in the hymenium, projecting or not, 60-70 x 4-7.5 um. Basidia
clavate, 4-sterigmate, becoming very thick-walled and vacuolate at
maturity, with a basal clamp, 36-50 x 7.5-9 um. Basidiospores ellip-
soid, hyaline, smooth, negative in Melzer’s reagent, with a single
large guttule, (6-)8-11 x 4.5-6.5 um. Comments - The specimen
illustrated had a number of spore forms present which showed various
degrees of distortion. Voucher specimen - JPL 1224.
CERACEOMYCES SERPENS (Fr.) Ginns, Can. J. Bot. 54:147. 1976.
Basidiocarps effused, shallow-merulioid to reticulate-poroid, remaining
so upon drying, hymenium separable, yellowish (2.5Y 8/4) to pinkish
526
Vg > ae Na es Armillaria mellea (JPL 1224). a) subhymenial
hyphae; b) tramal hyphae; c) sterile hymenial elements; d)
basidia, most with thickened walls and large vacuoles; e)
basidiospores; f£) distorted basidiospores.
buff (2.5YR 8/2 to 5YR 8/4); margin smooth, white, broad, fimbriate,
weakly rhizomorphic. Hyphal system monomitic; subicular hyphae
thin-walled, nodose-septate, occasionally branched, 2-6 um diam.
Sterile hymenial elements lacking. Basidia cylindric-clavate, 4-ster-
igmate, with a basal clamp, 28-32 x 5.5-7 um. Basidiospores hyaline,
smooth, short-cylindric, negative in Melzer’s reagent, 4-5 x 2-2.5um.
Voucher specimen - JPL 1309.
CERATOBASIDIUM CORNIGERUM (Bourd.) Rogers, Univ. Iowa Studies 17:5.
1935),
Basidiocarps very thin, shiny gray, adherent, a smooth film on the
substratum; margin indistinct, abrupt. Hyphal system monomitic;
subicular hyphae thin- to slightly thick-walled, simple-septate, often
collapsing in the subhymenium and difficult to make out, 5-5.5 um
diam. Sterile hymenial elements lacking. Basidia subglobose to
short-clavate, with four broad sterigmata, simple-septate at the base,
13-15 x 6.5-8 um. Basidiospores hyaline, smooth, ellipsoid, negative
in Melzer’s reagent, strongly cyanophilous, a very few germinating by
repetition, 6.5-8 x 4-5 um. Comments - This collection was made on
charred oak wood. Only immature basidia with short sterigmata were
observed. Voucher specimen - JPL 1077.
GLOEOPHYLLUM SEPIARIUM (Wulf.:Fr.) Karst., Finl. Hattsv. 2:80. 1879.
Basidiocarps reflexed, sessile, tough-leathery, usually in scattered
groups on slash; pileus' surface felty to matted-tomentose to strigose
rat
Fig. 2. Ceraceomyces serpens (JPL 1309). a) hypha from base
of basidial candelabrum; b) subicular hyphae; c) basidia; d)
basidiospores.
Fig. 3. Ceratobasidium cornigerum (JPL 1077). a) subicular
hyphae; b) basidia; c) basidiospores.
with concentric zonation in older specimens, the darker areas umber
(5YR 2/2), grading to rusty (near 5YR 5/8) and tan (near 10YR 8/6);
hymenial surface radially lamellate, grading to daedaloid in older
regions, concolorous with the lighter areas of the pileus. Hyphal
system dimitic; generative hyphae hyaline, thin-walled, nodose-septate,
2-2.5 um diam; skeletal hyphae brown, aseptate, thick-walled, rarely
branched, 2.5-4 wm diam. Cystidia hyphoid, not projecting, thin-
328 Lindsey - 4
walled, tapering to an obtuse apex, 34-38 x 4-4.5 um. Basidia slender,
cylindric-clavate, 4-sterigmate, varying in length, with a basal clamp,
17-66 x 4.5-6 um. Basidiospores hyaline, smooth, thin-walled, cylin-
dric to slightly curved, negative in Melzer’s reagent, 7.5-10 x 2.5-3.5
um. Comments - this collection represents the second brown-rot fungus
collected on Gambel oak in this series of papers. G. sepiarium is
common on conifers but but rare on Gambel oak. The other specimen was
Gloeophyllum trabeum (Lindsey, 1985). Voucher specimen - JPL 1373.
Fig. 4. Gloeophyllum sepiarium (JPL 1373). a) nodose-
septate generative hyphae; b) skeletal hyphae; c) cystidia;
d) basidia; e) basidiospores.
HYPHODERMA ANASAZIENSE Linds., sp. nov.
Basidiocarpi annui, ochracei, effusi-resupinati; hymenii superficies
laevis, coacta. Hyphae septatae, fibulatae, 3-5 wm diam; hyphis
fasciculis adsunt in subiculo. Cystidia nulla. Basidia tetraspora
clavata, fibulata, 28-36 (8 14.5=679. yim, Basidiosporae hyalinae,
tenuitunicatae, laeves, cylindraceae vel eellipsoideae, 5.5-7.5 x
3.0-4.0 um. HOLOTYPUS: J.P. Lindsey 1297, on Quercus gambelii
Nutt. slash, 8/2/84. East Animas Road, LaPlata County, CO, in herb.
National Fungus Collections, Beltsville, MD, U.S.A. (BPI); isotype in
herb. ARIZ.
Basidiocarps annual, effused-resupinate, pellicular, felty, smooth,
ochre-yellow (near 10YR 8/6 to 2.5Y 9/4); subiculum more loosely
arranged, with conspicuous cordons near the substrate, concolorous with
the hymenium; margin thinning out to abrupt, fertile. Hyphal system
monomitic; subicular hyphae hyaline, thin-walled, nodose-septate, 3-5
um diam; basement cordons present in deep subiculum, light golden, the
hyphae slightly rough-walled and nodose-septate. Sterile hymenial
Lindsey - 5 329
elements lacking. Basidia long-clavate, 4-sterigmate, with a basal
clamp, 28-38 x 4.5-6.5 um. Basidiospores hyaline, smooth, negative in
Melzer’s reagent, cyanophilous, cylindric-ellipsoid, 5.5-7.5 x 3.0-4.0
um. Voucher specimen - JPL 1297 (HOLOTYPE)
Fig. (5) Hyphoderma anasaziense CIPEVT A297): a) nodose-
septate subicular hyphae; b) basidia; c) basidiospores; d)
nodose-septate hyphae from subicular cordons.
HYPHODERMA MEDIOBURIENSE (Burt) Donk, Fungus 27:15. 1957.
Basidiocarps on well-decayed wood, smooth to slightly tuberculate,
effused, buff to pinkish-buff, (10YR 9/2 to 7.5YR 8/2), cracking on
5.20
Fig. 6. Hyphoderma medioburiense (JPL 1390). a) subicular
hyphae; b) leptocystidia; c) basidia, showing oily contents;
d) basidiospores, with abundant oily contents.
drying, appearing slightly punctate under higher magnifications (30x)
due to resinous accumulations at the cystidial apices; margin abrupt to
ayia |
abruptly thinning out. Hyphal system monomitic; subicular hyphae
thin-walled, nodose-septate, 3.5-4.5 um diam. Leptocystidia clavate to
long-sinuous, some projecting up to 30 um, others enclosed, thin-
walled, with a basal clamp, on the average up to 95 (but some up to
165)". x. 7-10 um. Basidia clavate, 4-sterigmate, with oily contents,
with a basal clamp, 32-40 x 6-9 um. Basidiospores hyaline, smooth,
cylindric to allantoid, negative in Melzer’s reagent, with abundant
oily contents, 11-14 x 4-6 um. Voucher specimen - JPL 1390.
KAVINIA HIMANTIA (Schw.) John Erikss., Sym. Bot. Upsal. XVI:1, p. 160.
1958.
Basidiocarps loosely adherent, resupinate, becoming broadly effused;
fertile area byssoid, hydnaceous, the spines up to 0.5 mm long, buff
colored (10YR 9/2); margin white, arachnoid to weakly fimbriate, with
abundant buff to white rhizomorphs, these penetrating the wood and
surrounding humus. Hyphal system monomitic; subicular hyphae hyaline,
smooth to roughened, some with coarse crystalline encrustation, some
becoming slightly thick-walled, some ampullate at the septa, 1.5-3.5ym
diam. Basidia clavate, 4-sterigmate, some pleurobasidia present,
developing along edges of teeth and growing outward, variable in
length, with a basal clamp, 30-50 x 5-6 um. Basidiospores slightly
olive-green, smooth to bumpy, (the irregular outline seen most readily
in Melzer’s reagent), negative in Melzer’s reagent, narrowly ellipsoid
to subfusiform, 8.5-10 x 3-3.5 um. Voucher specimen - JPL 1371.
Fag. 3). Kavinia himantia (JPL 13/71). a) thin-walled
subicular hyphae; b) thick-walled subicular hypha, showing
ampullate septa; c) basidia of varying lengths; d) smooth to
bumpy basidiospores.
PHANEROCHAETE SANGUINEA (Fr.) Pouz., Ceska Mykol. 27:26. 1973.
Basidiocarps adnate, smooth, effused in small patches, yellowish-tan
with light orange areas (10YR 9/4 to 7.5YR 8/4), not cracking, orange
areas green in KOH; margin strongly fimbriate, white, with occasional
pale yellowish rhizomorphs. Hyphal system monomitic; subiculum a
"textura intricata" (Burdsall, 1985); subicular hyphae thin-walled and
richly branched in the subhymenium to thick-walled near the substrate,
mostly simple-septate with occasional single and double clamps in the
deep subiculum, 3.5-6(-8) wm diam. Cystidia abundant, thin-walled,
tapering to an acute apex, simple-septate at the base, 46-58.5 x 4-5
um. Basidia slender-clavate, 4-sterigmate, with a simple septum at the
4
base, 33-39 x 3.5-5 um. Basidiospores cylindric-ellipsoid, hyaline,
smooth, negative in Melzer’s reagent, often with one or two guttules,
4-5.5 x 2-2.5 um. Voucher specimen - JPL 1374.
Fig. G. Phanerochaete sanguinea (JPL 1374). a) nodose-
septate hyphae from deep subiculum; b) simple-septate
subicular hyphae of varying wall thickness; c) leptocystidia;
d) basidia; e) basidiospores.
PHANEROCHAETE SORDIDA (Karst.) John Erikss. et Ryv., in John Erikss.
et al., Cort. of North Burope; 1521023." 1978
Basidiocarps effused on bark, pinkish-buff (10YR 9/4), smooth, compact,
membranaceous, cracking on drying to reveal a loose, white subiculum;
margin thinning out, white, farinaceous to finely fibrillose under a
10X lens. Hyphal system monomitic; subicular hyphae simple-septate,
thin-walled in the hymenial area, 2-2.5 wm diam, thick-walled in the
deeper areas of the subiculum, 3-7.5 um diam; no double or multiple
clamps seen. Cystidia few, cylindric, not encrusted, thin-walled at
apex to thick-walled at the base, some with a short lateral projection
near the apical end, 52-98 x 4.5-7 um. Basidia clavate, 4-sterigmate,
Sa0
Fig. 9. Phanerochaete sordida (JPL 1221). a) thin-walled
subcandelabral hypha; b) thick-walled subicular hyphae; c)
leptocystidia; d) basidia; e) basidiospores.
with a simple septum at the base, 19-26 x 3.5-4 um. Basidiospores
cylindric-ellipsoid, hyaline, smooth, negative in Melzer’s reagent,
(4-) 4.5-6 x 2-3 um. Comments - Eriksson et al. (1978) and Burdsall
(1985) discuss the extreme variability of this species. The sole
collection examined in this paper had no encrustation on the cystidia
and the cystidia were highly variable in length, with most being
short. Only two cystidia were seen that possessed the lateral branches
discussed by Eriksson et al. Voucher specimen - JPL 1221.
PLATYGLORA LAPLATA Linds., sp. nov.
Basidiocarpi albidi, farinacei, byssoidei, laeves vel irregulariter
verrucati, margo abruptus vel fimbriatus; basidiocarpi parasiticus in
basidiocarpo Peniophora nuda. Hyphae septatae, fibulatae, 3-4.5 um
diam. Cystidia nulla. Basidia cylindracea, tetracellularia, 90-106 x
8-10 um; sterigmata vel 14 x 3 um; basidiosporae hyalinae, laeves,
tenuitunicatae, ovoideae, germinans iterum adque iterum, 11.5-17 x
7.5-13 um. HOLOTYPUS: J.P. Lindsey 1279, on Quercus gambelii
Nutt. slash, 9/29/84. Sheep Springs drainage, LaPlata County, CO, in
herb. National Fungus Collections, Beltsville, MD, U.S.A. (BPI);
isotype in herb. ARIZ.
334
Basidiocarps whitish, byssoid, farinaceous, smooth to irregularly
warted, growing adjacent to and _ touching a healthy Peniophora nuda
basidiocarp; margin abrupt to finely fimbriate Hyphal system monomi-
tic; subicular hyphae conspicuously nodose-septate, thin-walled, 3-4.5
um diam. Sterile hymenial elements lacking. Basidia long-cylindric,
arising directly. from thin-walled hyphae, 4-celled, usually with a
basal clamp, 90-106 x 8-10 um; sterigmata up to 14 x 3 um, arising from
the upper end of each cell. Basidiospores hyaline, smooth, ovoid,
negative in Melzer’s reagent, with a blunt apiculus, germinating by
repetition laterally or opposite the apiculus, 11.5-15(-17) x 7.5-11
(-13)um. Comments - although this specimen was touching a collection
of Peniophora nuda, and presumably parasitizing it, it clearly does not
fit the existing spore descriptions of Platygloea peniophorae Bourd. et
Galz. as discussed by Martin (1940). The spores of my collection are
much larger than any reports found for P. peniophorae. Voucher
specimen - JPL 1279.
Fig. 10. Platygloea laplata (JPL 1279). a) nodose-septate
subicular hyphae; b) basidia; c) basidiospores, some germin-
ating.
Bee
SUBULICYSTIDIUM LONGISPORUM (Pat.) Parm., Consp. Syst. Cort. p. 121.
1968.
Basidiocarps thin, farinaceous, patchy, white to dingy; margin indis-
tinct, concolorous. Hyphal system monomitic; subicular hyphae thin- to
slightly thick-walled, nodose-septate, often encrusted with plate-like
crystals similar to those on the cystidia, 2.5-3 um diam. Cystidia
abundant, slender, thick-walled, subulate, heavily encrusted with
flattened, evenly spaced crystals except at the extreme apex, with a
basal clamp, 45-66 x 2.5-4.5 um. Basidia clavate, 4-sterigmate, some
showing a median constriction, with a basal clamp, 11.5-13.5 x 3.5-4
um. Basidiospores hyaline, smooth, acicular, somewhat flexuous,
negative in Melzer’s reagent, usually with several uniseriate guttules,
(12-) 13.5-22 x 1-1.5 um. Voucher specimen - JPL 1222.
Big. iis Subulicystidium longisporum (JPL 1222). a)
nodose-septate subicular hyphae, some encrusted; b) encrusted
cystidia; c) basidia; d) basidiospores.
TOMENTELLA MOLYBDAEA Bourd. et Galz., Bull. Soc. Myc. France 40:142.
1924.
Basidiocarps effused-resupinate, granulose, pinkish-buff (near 10R
7/2); margin thinning out, grayish. Hyphal system monomitic; subhymen-
ial hyphae thin-walled, nodose-septate, hyaline, turning light bluish-
black in KOH; subicular hyphae hyaline, thin- to slightly thick-walled,
nodose-septate, 3-5 wm diam; cordons lacking. Sterile hymenial
elements’ lacking. Basidia clavate, 4-sterigmate, with granular
contents staining bluish-black in KOH, with a basal clamp, 38.5-57 x
6.5-9 um. Basidiospores light brown, echinulate, negative in Melzer’s
reagent, globose to irregularly globose to lobed, 5.5-7 x 5-5.5 um.
Voucher specimen - JPL 1375.
Fige ize Tomentella molybdaea (JPL 1375). a) subicular
hyphae; b) basidia; c) basidiospores.
TOMENTELLA SUBTESTACEA Bourd. et Galz., Bull. Soc. Myc. France
M02144. 1924.
Basidiocarps pale buff (7.5YR 7/2), mucedinoid, arachnoid to granulose,
effused on well-decayed wood; margin thinning out, grayish white.
Hyphal system monomitic; subicular hyphae hyaline, slightly thick-
walled, abundantly branched and nodose-septate, 3-4 wm diam. Cystidia
thin-walled, obclavate to capitate, 21.5-58 x 4.5-8 um, the capitate
apices 3.5-5 um diam, with a basal clamp, some with one or two clamp
connections along their length. Basidia long-clavate to cylindric,
4-sterigmate, some with intercalary septa, with a basal clamp, 38.5-56
x 7.5-10 wm. Basidiospores dull brown, echinulate, globose to lobed,
negative in Melzer’s reagent, 6-7 x 4.5-6 um. Voucher specimen - JPL
1389.
TRECHISPORA FARINACEA (Pers.:Fr.) Liberta, Taxon 15:318. 1966.
Basidiocarps effused in small patches, farinaceous to grandinioid,
white to grayish-yellow (5Y 8.5/1 to 7.5Y 8/2); margin white, arach-
noid, weakly rhizomorphic. Hyphal system monomitic; subicular hyphae
thin-walled, nodose-septate, some ampullate at the septa, 2-5.5 um
diam. Sterile hymenial elements lacking. Basidia short-cylindric to
clavate, urniform at first, 4-sterigmate, with a basal clamp, 10-13.5 x
4-5 um. Basidiospores hyaline, finely echinulate, ovoid, negative in
Melzer’s reagent, 3-4 x 2-3 um. Voucher specimen - JPL 1367.
Sod
Fig. 13. Tomentella subtestacea (JPL 1389). a) subicular
hyphae; b) cystidia; c) basidia; d) basidiospores.
QUQGE
ics
§ O96 eS
Fig. 14. Trechispora farinacea (JPL 1367). a) nodose-
septate subicular hyphae with ampullate septa; b) basidia; c)
basidiospores.
TRICHOLOMOPSIS PLATYPHYLLA (Fr.) Sing., Schweiz. Zeitschr. Pilzk.
T5963. 1939.
Basidiocarps centrally stipitate, single to scattered on large slash;
pileus 3.5-7 cm diam, flat to slightly depressed, shiny gray-brown
(near 10YR 7/4), fibrillose; context thin, white; lamellae white to
338
cream, adnate to adnexed, distant; stipe 0.5-1 cm diam, equal or
slightly enlarging at base, smooth, twisted, showing spiral striations;
annulus lacking. Hyphal system monomitic; tramal hyphae of two types:
one type thin-walled, nodose-septate, the clamps often distorted,
2.5-8.5 wm diam; the second type inflated, slightly thick-walled,
aseptate, tapering to a blunt apex, up to 22 um diam. Pleurocystidia
hyphoid to clavate, not projecting, thin-walled, with a basal clamp,
40-45 x 6 um; Cheilocystidia thinner, similar. Basidia clavate, with a
narrowly tapered base subtended by a clamp, with abundant oily con-
tents, 39-41 x 8-9 um. Basidiospores hyaline, smooth, ellipsoid to
ovoid, negative in Melzer’s reagent, 8.5-10 x 5.5-7.5 um. Voucher
specimen - JPL 1082.
Fig. 15. Tricholomopsis platyphylla (JPL 1082). a) subhy-
menial hyphae; b) tramal hyphae of various diameters, some
apparently aseptate; Cc) pleurocystidia; d) basidia; e)
basidiospores.
PHAROMARASMIUS ERINACEUS (Fr.) Kiihner. Encyc. Myc. 12331 tae
Basidiocarps reviving when moistened, up to 1 cm broad and 1.5 cm high,
on attached dead branches or slash; pileus depressed, roughly scaly-
hairy, near 2.5YR 3/4; stipe central, concolorous with pileus, equal,
wooly; annulus none; lamellae white to pinkish. Hyphal system mono-
mitic; tramal hyphae hyaline, thin-walled, with clamps, 3-4 um diam;
Sog
hyphae comprising pileus tomentum pseudoamyloid in Melzer’s reagent,
corrugated, pigmented, thick-walled, with clamps, 4-8 um diam.
Cystidia of two types: pleurocystidia fusoid, 30-42 x 7-10 um,
sometimes absent; cheilocystidia fusoid, 20-35 x 4-9 um. Basidia
clavate, 4-sterigmate, with a basal clamp, 35-43 x 7-8 um.
Basidiospores ovoid to short-ellipsoid, lightly pigmented,
straw-colored in KOH, smooth, negative in Melzer’s reagent, 8-10 x
4.5-6 wm. Comments - Smith and Hesler (1968) call this species
Pholiota erinacea (Fr.) Reas The light pinkish spore print, however,
led Singer (1963) to refer it to Phaeomarasmius. Personal communica-
tions with Dr. Roy Halling and Dr. David Largent suggest that it is
best left in this genus. This fungus is not uncommon on slash and dead
attached limbs. It causes a stringy white rot. Voucher specimens -
JRE L5is FI PELS1L190; J Pil 368"
Fig. 16. Phaeomarasmius erinaceus (JPL 1151). a) subhymen-
ial hypha; b) encrusited hyphae from pileus tomentum; c) hypha
from trama of pileus; d) basidia; e) basidial candelabrum; f)
basidiospores.
CONCLUSIONS
Sixty-six species of wood-inhabiting basidiomycetes have been
reported on Gambel oak from southwestern Colorado in a series of three
papers. These fungi play differing roles in the ecology of the host
tree. Most are active decomposers of slash on the ground. The most
conspicuous of these are Polyporus arcularius and Pycnoporus cinnabar-
inus. Some fungi are most frequently associated with dead branches on
living trees. Exidia glandulosa and Hyphodontia crustosa are predom-
inant in this category. Finally, Inonotus andersonii is a parasite of
living trees; it kills out tops or sometimes entire trees. iieas
extremely common in the clonal scrub stands of oak in this area.
Most of the fungi identified on Gambel oak are white-rot fungi.
Brown-rot fungi are common on associated trees, such as Pinus ponderosa
340
Laws. and Populus tremuloides Michx. It seems likely that brown-rot
fungi do not favor the oak substratum. This is of interest in relation
to the work of Jurgensen et al. (1977) which points out the importance
of brown-rot fungi in soil building, mycorrhizal formation, and water
retention. Many fungi not reported in this series of papers ultimately
will be (and have been) found on this substratum, especially in areas
such as Utah, Arizona, and New Mexico, where environmental conditions
favor different kinds of fungi. However, the sixty-six species
reported seem to represent a fair cross-section of the mycota of this
area. These wood-rotting fungi are important in returning organic
materials to the soil. Many oak brush stands are on extremely steep
slopes, where they stabilize the soil against mud slides and less
severe forms of erosion. The white rot fungi predominating on oak
contribute, however slowly, to the soil building process. Because of
the continued interest in eradication of Gambel oak through burning and
herbicide treatment for improvement of wildlife grazing and cattle
pasture, and because of the increased interest in this species for
firewood, it is critical that we know more about the ecology of the oak
woodlands in this area. Further studies on the impact of long-term oak
brush eradication should be pursued.
KEY TO SPECIES OF WOOD-ROTTING BASIDIOMYCETES
THAT DECAY GAMBEL OAK
la. Basidia of the holobasidium type ............... g's mwa te oe Late at acta
lb.: Basidia ofthe heterobasidium ity pes’. 0. is. eines sy es «sins Prior. y.
La eu DASTOLANDD LUL CATON cit ojsaiels als « wc -ate'w alee alelelele ts Dacrymyces minor
2b. Basidia cruciately septate or transversely septate ........ 3
3a. Basidia cruciately septate; basidiocarp gelatinous,
Dl aACK LO) PLOW WG ee tateieus Cle ela Scieiaala Bic sip tate ln cere ated Exidia glandulosa
3b. Basidia transversely septate; basidiocarp
AL Wp WEREMIGs, Wessel cte la iaia''s tere gaa ae ieniieiattue alee aie .+eeee- Platygloea laplata
4a. Basidiocarp agaricoid, fleshy, with hymenium
borne on radiating lamellae ...........6.. Aides ete Ae eT
4b. Basidiocarp poroid, toothed, smooth, or merulioid; if
fleshy, not with hymenium borne on radiating lamellae;
if ‘lamellare, then’ tough.and Librous:. . v8. sss 000s ee ema reece
5a. Basidiocarps eccentrically stipitate to sessile ..... eetetetntate opal we
5b.) (Basidiocarps centrally stipitate ..5.40. 2. « 60s hat one: erate ura: eerie Ae:
6a. Basidiocarps minute (1-3 mm diam), bluish-black;
basidiospores globose ......... Sisko: shane Resupinatus applicatus
6b. Basidiocarps larger, buff-colored; basidiospores
OMOL Git On SROL © OV U1 IVER LCi etal scence locate oi cloes leipternce 1 aie site. a stele aseeneenl
7a. Basidiospores minutely echinulate, amyloid
Li MOLZer? SareCAren ty li. la waipiele + visto lores vie alate Lentinellus cochleatus
7b. Basidiospores smooth, negative in Melzer’s reagent ... Panus rudis
8a. Spore print pure white, basidiospores smooth ........++++-- 10
8b. Spore print pinkish, clay-colored or brown, spores smooth
OTIC UBOE CULE t Cie cim n'ls sbels tele tsi cl above ose ay senuedple sh asstale etvcainvaly chelator sin, ame
9a.
9b.
lay
11b.
i3a.
13b.
Tar
15b.
17a.
17D:
19a.
19b.
Zia
21b.
341
Basidiocarps orange to rust-colored; basidiospores
ellipsoid to broadly fusiform, verrucose,
6.9-9 % 4.5-6 pms .n es RPsos SAvabenoteis a ea thre) ates ave Se tore Gymnopilus sapineus
Basidiocarps small, brown, hairy; basidiospores ovoid to short-
BLT pS010, 36-10 xX) 4s. —O GMM ve sie as oon sls eels « Phaeomarasmius erinaceus
10a. Annulus present; young pileus surface with fine,
POTINUCOMSCA LOS) ctew acts ems wins s shale cw lee ass Ue Wie Armillaria mellea
10b. Annulus lacking; pileus surface glabrous to
ADPLESSEd TADTILLOS@ ewe cle ccs 6's Tricholomopsis platyphylla
Basidiocarp tough, not fleshy; hymenophore poroid to
USCGSLOV OL) LAGTALLY | LAMCMLATO( 60 5012's 5's boa cetele le Goleta lee hele eure le en We
Basidiocarp not fleshy; hymenophore hydnoid,
MELD IZ OLA UT SMOD ED MOL AE CeEQULAE siete tse levers les wees ee relshelela se aera sieeve 26
eae Y DUA Le Sy SCCM MONOML TLC! i524 ts cle oic1e'o > sft o/c snc e es lerare Wels dielale™® 06 13
LZORE Hy DNaleSVSTCMPOLer OG CL IMLtaCs 4 .ccne ss Ge aie a aleve eeve see wialele evel's 18
Generative hyphae nodose-septate ......... sarah train ata te Lie tee: Gis ereRs wae
Generative hyphae simple-septate ..... RNC ae AIS ry sete cr de er 15
14a. Basidiocarps effused-resupinate,
CrYEAM=COLOECC! cis sc icicles ori els eipie let ers ooh Ceriporiopsis mucida
14b. Basidiocarps resupinate to pileate,
reddish-brown with a golden pore surface,
COT CHErLy Lear ahi KONs 5 oss eo aierare eee Hapalopilus nidulans
Basidiocarps resupinate, whitish to pinkish-tan ..............- 16
Basidiocarps hoof-shaped, golden-brown, with a
EGIL COLE. uk cls cs er eiera's 5 ae ei lela G15 Niels aiWils!s later’ Inonotus dryophilus
16a. Basidiospores reniform to
Broad LY allan told aie <sics.¢ sere atostec's-cie & slat Ceriporia reticulata
i6b. Basidiospores cy lindric=elli psoidi. 2)... \cje ce s)he we wale oo nie
Basta OSD0Lres 42450 ex 2-255) UM, mises ee Vad sok eee etele Ceriporia tarda
Basal 0Spores 4X7 3-355" UR: 5 2% ais Sieleerece ere o's ta ele Ceriporia excelsa
LOAveHY PRALPESYS COMM EL AMT ULCuls ess cs dae Go a, oes ale C16) %oiiele elie 0s exer! ele 19
18b. Hyphal) system dimitie. >. ..2../:.- Bhelaterehe so a edeie ies ame elet oe tote rena re ts ae
BAStaOCAar pS ew LCS aise pav-5 01518 Wek als nis 46s bo heieia lt orca tale aly wie eros 20
Basidiocarps bDraghtvorange) uss 6 te. ss ac Pycnoporus cinnabarinus
20a. Basidiospores hyaline, thick-walled,
dextrinoid in Melzer’s reagent,
Dee o ee. as OT LIMO ly a os le sa cis 6 es ale 68 Perenniporia medulla-panis
20b. Basidiospores hyaline, allantoid, negative in
Melzer’s reagent 3.5-5 x 0.5-1 um ....... Incrustoporia nivea
Basidiocarps brownish-grey to blackish ..........+... hel se. wtoletnts % Ze
Basidiocarps tan to bright-colored ...........6.. wm eheta tas Rigre ae S06 aa
22a. Hymenophore ranging from poroid to
Gaedaloidata Jamel later su s's.0 5 «aisles + o's 0.8 os «ss sie clelease se eo sis 6 23
22b. Hymenophore distinctly poroid .......ccsccccsesccesvcescas 24
342
Leave
23b.
25a.
25b.
27as
27d.
29a.
29b.
31a.
31b.
33a.
33b.
25a,
35b.
Basidiospores 7.5-8.5 x 3.5-4.5 um; basidia up to
ZOE {MAL ONG Sipe saat eie, <l'si shaseibyets phone lee oiwln ene scene eee ete Gloeophyllum trabeum
Basidiospores 7.5-10 x 2.5-3.5 um; basidia up to
GG Fpmpl on Paras: Ria cients, 0 sever sts oeke ves eters ne cates Gloeophyllum sepiarium
24a. Basidiospores pigmented; basidiocarps fruiting
between) bark andgwood) octc5 5%). tesevcidi ete wena Inonotus andersonii
24b. Basidiospores hyaline; basidiocarps fruiting on
GEdd IDVARCNES wars oison hale Theiss eles wate outels Phellinus ferruginosus
Basidiocarps tan; centrally stipitate ....... Polyporus arcularius
Basidiocarps white to bright lemon-yellow;
CLTUSEdSYeESUPINALS i501. sieloteis.s «ve eis « prepelie te oases eke Perenniporia tenuis
26a. Basidiocarps with a distinctly loose to densely
toothed hymenophore ........... ale guste Sus vere aha hele siete oe eT cacy, i)
26b. Basidiocarps with hymenophore tuberculate, smooth,
merulioid, or, otherwise? ccs sss \v7eid. saree eiaie ae neon te ete tees 32
Basidiocarps densely aculeate to hydnaceous ............ sifo'skeitte RCO
Basidiocarps, wartedto furfuraceouss ss:a5~ sowie seo eS es Sy
28a. Basidiocarps white to cream colored or buff .............. 29
28b. Basidiocarps vinaceous; margin strongly fimbriate
and ThiZomorphic 4% nw «28 sais ote Steccherinum fimbriatum
Lagenocystidia present is a0 ssie «stale «is o.ofete 2 wes sure Hyphodontia arguta
Lagenocystidias Lack ine: :.5 2 sm eles: stage ie s/sta d sleielat eg delnlers io sla sce een 30
30a. Basidiocarps loosely attached, hydnaceous; margin white,
Wi thea Dundane. Fig ZOMOT DNS) seu os asic ep chs tole aie Kavinia himantia
30b. Basidiocarps strongly adherent, aculeate, with fimbriate
apices to the aculei ........ eeeeeeeee. Hyphodontia crustosa
Basidiocarps waxy, irregularly warted to folded
to hydnoid ..... Reayeh aks oracle ia'e a iscsi chels eraiszs eta soho siete eceie ts Phlebia albida
Basidiocarps porose-reticulate to furfuraceous,
in placesywith «blunt aculeiy.:. aotoi site wisn. Hyphodontia subalutacea
32a., Basidiocarps#felty tovarachhoidsto-byssoidy... ieee ee eee 33
32b. Basidiocarps pellicular to compact ....... a) owas Je «piste aman
Cystidia gpresemieier so esi. tls ots sis's id cats @ ais eee bates wc ceiane vias alts! Bevele’ emeee
Cystidia and other sterile hymenial elements lacking .......... 36
34a. Cystidia projecting, hyphoid, abundantly clamped,
lightly encrusted; basidiopores hyaline,
SMOO UM Miao ties shel esse nice glo icla! Hie alo! os Suatal ooactadenohe Amphinema byssoides
34b. Cystidia obclavate to eoedy capitate; basidiospores
pigmented, rough—-walleds .% . ./c/.1 sls + selewlajars ois a ialer A raireryy i ctie ee.
Basidiocarps pale buff; cystidia obclavate to capitate,
the capitate apex up to 5 um diam ......... Tomentella subtestacea
Basidiocarps gray-brown; cystidia golden, broadly capitate,
the apical cel) up cto) i4-umediam <2 .)taccs, ses! Tomentella pilosa
Biviale
37 bs
39a.
39b.
4la.
41b.
43a.
43b.
45a.
45b.
47a.
47b.
49a.
49b.
343
36a. No basidia present in hymenium; thick-walled conidiophores
and limoniform conidia present ............. Oidium candicans
36b. Basidia present in hymenium ..... Ris: l eer eo ieee a eer a ate 4 ios Pest,
Basiaiocarps darkareddisnabrownetOl rus te. %. ~ cee. vesstes mess oe se 38
BasidaGcarpseburc., WHILE. uCLAYi, OF CVCLLOWS «ccs specie ite vie fo. cies es 39
38a. Basidiocarps rust-colored; basidiospores lobed or
TELECULAL, sO (Xe) SO MUIM ec trace ciate slain es Tomentella ferruginea
38b. Basidiocarps dark reddish-umber; basidiospores globose,
aculeate,)/.5=9 [Umcdiam ... ss egs esis Tomentella ramossissimum
Basidiocarps with a distinct pinkish-buff
SE UU dateta tals for atal stole cise a iate eins. acbosis: se ia'e ei ei 6 on er ehekel o Tomentella molybdaea
Basidiocar psawnatish. VOLLOWwisn 1. COnDEOWNLSI i cclere 1s si eleis le srerere sie 40
40a. Basidiospores smooth, limoniforn,
OLADLCULAT GC icc Wars semanas sisteloretniercs or eiic Botryobasidium botryosum
40b. Basidiospores echinulate, ovoid to ellipsoid ............. 41
Basidiocarps whitish, farinaceous to grandinioid;
bas idposporesis—4eXe2—3 UM o5 o.5;0 «0's 5.0 cio se Besa Trechispora farinacea
Basidiocarps separable, olive to dull yellow, felty, with abundant
cordonsssbasidiospores 4.5-5 x 3=3.5.. UM vesicecdes Trechispora vaga
42a. Sterile hymenial elements (setae, acanthophyses,
CYSTIGLa neeTC 1) eLACRAMNG tereaia cunssiaie oe aber ohei ee Shar ehe tr els ane eueteneres ates 43
4a2b. sterile hymenial elements presen hives. c,d aid se tie whe aic.s Seale 9 sil
Basidiocarps pelliciular, separable, not felty 0.0.0.0 05.6000 44
Basidiocarps attached, adherent: aij ciiais «cee sinvice 5<< a Gur A 6 ae 46
44a. Basidiospores globose to subglobose ...... Athelia coprophila
BaD RED AS HULOSDOLES FC. SOU cs aie sie cies tei ays sl erei.caseveieisis foie wba oeaealela sis 45
Basidiospores narrowly ellipsoid to
De soa BOM gia ON a2 ay RENO 1) 3) oo) ahs ds) ots) her ec.stel ole iarcb alta Athelia bombycina
Basidiospores ellipsoid,
ED Se OUEK = Crt LIM, Lace ode siete tks State ae ta eles eyaler oie « Fibulomyces mutabilis
46a. Basidia short to slender-clavate, 4-sterigmate ........... 47
46b. Basidia utriform, (4-)6-8 sterigmate...Sistotrema brinkmannii
Basidiocarps shallow-merulioid, reticulate-poroid or
slightly phlebioid (hand lens may be necessary) ......eeeeeeees 48
Basig1Ocarps SMOOTNE LO. UDELCULAEC ya... cle oc cleis «oie isie « sisieielele © sleretelels « 49
48a. Hymenium shallow-merulioid to reticulate-poroid, yellowish to
pinkish-buff£; weakly rhizomorphic ...... Ceraceomyces serpens
48b. Hymenium smooth to slightly phlebioid, cracking along folds
upon drying, pinkish-buff to pinkish-gray; rhizomorphs
present, penetrating into wood and bark ... Phlebia lilascens
Basidiocarp a shiny, thin film on the
BUDS LLG UG eri sas! stecieaciie Alec's, « aisitie eae @ae 8 acess Ceratobasidium cornigerum
Basidiocarp thicker, subiculum often visible (with hand
lens) through cracks or pock-marks in the hymenium ............ 50
344
HSilac
5ib:
Sais
53b.
55a
Dab,
Si/ae
Ss
59a.
59b.
50a. Basidiocarps white to cream or ochraceous, often cracking on
drying; margin white, fibrillose, rhizomophic; clamp
GOMMGC LIONS) LACKING aie ole oeiere sii teraiee . Phanerochaete tuberculata
50b. Basidiocarps felty-pellicular, yellow-ochraceous; margin
thinning out to abrupt; clamp connections
PESO Ge o5ie seis; wiaters Glare stele oie ore ein eitierene Hyphoderma anasaziense
Gloeocystidiaor pseudocystidia’ present! occa ichiaiscls felsic siete ee ae
Gloeocystidiayor pseudocystidia absent ). ..tsii's sce slcie cesses leslete Be bs
52a. Acanthohyphidia abundant; basidiospores amyloid in
Melzer! SUTeagen tt asmvascsie sisysleie sveleis totals Aleurodiscus cerrusatus
52b. Acanthohyphidia lacking; basidiospores negative in
NGL SErGSGreagen Fi eieis oc cists clove od wee aiaiee dia wie ohare tip aieoreta, a ehoudea epee
Basidiocarps grayish-blue
EOP YUN ULea se loracecoinietecne Bi Soh letay tl oe ec thahat ets ole b9 eehe Peniophora violaceolivida
Basidiocarps pinkish-buff to orangish-gray ....... eee ace eiteee 54
54a. Gloeocystidia 38-75 x 8-13 um; basidiospores
Gey Ce 9), UID isl ciel e-evnhevaieie ot mie eo Peniophora pseudoversicolor
54b. Gloeocystidia 42-48 x 8.5-11.5 um; basidiospores
DES AB OD (Rie te Sc OMA ese) domps ee mich saw eeaanel ae eten ee .... Peniophora nuda
Thick-walled pigmented or encrusted sterile hymenial
SLEMENUS |! PLESENG Gals sietdo o1.0:.01 0) o/shorsioim pier seatal Mone Biel share oraeete sims cake tally < Scoops
Sterile hymenial elements not thick-walled
CEXCEDU ATI DASE) eae so ais ew Sones Hisielislia deletes chaste mien pobre Se esting
56a. Darkly pigmented, subulate setae
present ...... ptuaun de alate ch ecetepeteyate Lncpayatine.cpehe laa Hymenochaete arida
56b. 1 Encrusted lamprocystidia present (.)<)..... <)ciscleelstoteloiwis sip oueetelemainolde
Cystidia amyloid, dissolving
BU 33h MEMS tera e sa sie iaiceliet es Be ies lana eet Ti cretates ia hatin Tubulicrinis calothrix
CYSTIGTAPROE AMY LOL Ae wheie leis: ois!ncccenters wipes Susans dha Spetena: scatineene Hatietadel ae
58a. Cystidia subulate, encrusted with flat, evenly-spaced
crystals except at extreme
ADOKy eheratets gf PANT ahasiets 16, hale ehaiel « tared eter eels Subulicystidium longisporum
58b. Cystidia large, projecting, thick-walled, heavily encrusted
at apex and up to 2/3 of their
length, 24iciee' ot aed shel alisdet ot cfele shales seat aieletetel ane Phanerochaete velutina
Basidiocarps dimitic; basidiospores amyloid in
Melzer’s reagent ......... vibe cin ble alee bios s at's oss, OLGLCUM IEE SUtIms
Basidiocarps monomitic; basidiospores negative in
Mel erleiereaeMt ie io.cssieye si his eletrpi alate ereralel ew ealstsieiete a Sievets hie [ose tee OU
60a. Basidiocarps yellow to orange ......-.eeeceees oleae. dco eats 2101
60b. Basidiocarps white to pinkish-buff .............. etieks ease at areas
345
6la. Basidiocarps yellowish-tan with light orange areas,
the orange areas turning green in KOH; cystidia thin-walled,
46-58.5 x 4-5 um; basidiospores
eS M IL = Dig JHU Ie circ ietalel shale esa itately »! 6 6\%6 ......- Phanerochaete sanguinea
61b. Basidiocarps bright orange- Gey ae when fresh, fading to a
dull yellow upon drying; cystidia thin-walled, 63-78 x
4.5-6.5 um; basidiospores
Ban =O) OX Lee STM cies e oie lak eieleiciels sss 8 e.5re 6 Phanerochaete carnosa
62a. Basidiocarps bright white; capitate, projecting cystidia and
embedded stephanocysts present ..... Hyphoderma praetermissum
G20. basidiocarps pinkish—white to DULLER 6 6. ous cide w sisi wares 6 63
63a. Leptocystidia thin-walled, long sinuous, some over
130 um long; basidiospores
Mee XA OV UM Palen swe oisls!e odie weiss) sine ol aio Hyphoderma medioburiense
63b. Leptocystidia thick-walled At base, mostly shorter than
PG0s1m; basidiospores ranging) 4-6 "x 2-3 Ym) os ie. awe oe asic anes 64
64a. Cystidia thin-walled,
SD SSO IMEX — 7a TIM ctetareer itis ere ssterens s.nce ele Phanerochaete arizonica
64b. Cystidia thin-walled at apex to thick-walled at base, some
with short lateral projections near the apical end,
LO OE Mane | BATE Leta hel ates tere are rer ali aicie tenes. eel ane Phanerochaete sordida
ACKNOWLEDGEMENTS
Thanks are gratefully extended to: Mrs. Donna Giersch for typing,
proofreading, and helpful suggestions; Dr. Roy Halling and Dr. David
Largent, for identification of Phaeomarasmius erinaceus; and Dr. R.L.
Gilbertson for critical review of the manuscript.
LITERATURE CITED
Burdsall, H.H., Jr. 1985. A contribution to the taxonomy of the genus
Phanerochaete (Corticiaceae, Aphyllophorales). J. Cramer, New York.
165"p.
Eriksson, John, Kurt Hjortstam, and Leif Ryvarden. 1978. The
Corticiaceae of North Europe. Vol. 5. Fungiflora, Oslo.
Jurgensen, M.F., M.J. Larsen, and A.E. Harvey. 1977. Effects of
timber harvesting on soil biology. pp. 244-260. In forests for
people. A challenge in world affairs. Proc. Soc. Am. Foresters.
Lindsey, J. Page. 1985. Basidiomycetes that decay Gambel oak in
southwestern Colorado. Mycotaxon 22:327-362.
Lindsey, J. Page. 1986. Basidiomycetes that decay Gambel oak in
southwestern Colorado: II. Mycotaxon 25:67-83.
Martin, G.W. 1940. Some heterobasidiomycetes from eastern Canada.
Mycologia 32:683-695.
Munsell Book of Color. 1976. (Ed.) Munsell Color Co., Inc., Baltimore
MD.
Smith, Alexander H. and L.R. Hesler. 1968. The North American Species
of Pholiota. Lubrecht and Cramer, New York. 394 p.
Singer, Rolf. 1963. The Agaricales in Modern Taxonomy, 2nd Ed.
Pholiota, pp. 550-558. Weinheim.
MYCOTAXON
Vol. XXVII, pp. 347-360 October-December 1986
Lahmia Kérber (= Parkerella A. Funk)
a misinterpreted genus with isolated position.
Ove Eriksson
Institute of Ecological Botany,
University of Umea, S-901 87 Umea, Sweden
ABSTRACT
The only species accepted in the genus Lahmia
Korber, L. kunzei Korber, has been considered
a non-lichenized discomycete with lecanor-
alean ascomata, but is shown here to differ
so much from other ascomycetes that it has to
be referred to a new family in a new order
(Lahmiaceae, Lahmiales). Parkerella Funk,
originally described as an aberrant genus in
Coronophorales, is a synonym.
INTRODUCTION
Some interesting fungi have been described on bark of
Populus spp. from North America during the two last de-
cades. Rough bark on P. balsamifera may be caused by the
bitunicate pyrenomycete Rhytidiella moriformis Zalasky
(1968: 1383, 1975: 780). The cork bark disease on P.
tremuloides is believed to be caused by another Rhytidiella
species, R. baranyayi Funk & Zalasky (1975: 752). Funk
(1976: 868) found that this species is often associated
with another fungus, which he described as Parkerella
populi gen. et sp. nov. in the order Coronophorales.
The first European record of R. moriformis was reported
on Populus tremula from N. Sweden a few years ago (Eriksson
1981: 51). This find initiated a search for aspens with
cork bark disease in Sweden. Rhytidiella baranyayi was
never encountered, but quite recently it suddenly occured
to me that Parkerella populi may be a fungus that has
been known from Europe for more than 120 years, viz. Lahmia
kunzei Kérber. This is a wide-spread and common, but over-
looked fungus on Populus tremula. It has been treated as a
348
discomycete, but it perfectly matches Funk’s description of
the pyrenomycete Parkerella populi. A closer examination
reveals that its morphology has been misinterpreted and
that it is not closely related to any order in current
classifications. The present paper gives information about
the nomenclature, morphology, systematic position, and
biology of this fungus.
MATERIALS AND METHODS
Semithin sections of material embedded in LKB 2218-500
Historesin were cut with glass knives in an LKB Histo-Range
Microtome. SEM graphs were made with a JEOL JSM-P15.
RESULTS
Lahmia Kérber 1861: 281.
Syn.: Parkerella A. Funk (non Parkerella Muniers-Chalmas ex
L. Morellet & J. Morellet, 1922; fossil alga; see
Farr & al. 1979: 1266), Can. J. Bot. 54: 686, 1976.
Lahmia kunzei Kérber 1861: 282
Syn.: Parkerella populi A. Funk, l.c.
Figs.: 1-3.
Ascomata scattered or occasionally 2 or 3 close together,
erumpent, turbinate, when dry with flat or slightly concave
upper surface (very young ascomata convex), swelling in
water and fertil portion then becoming subglobose and up
to ca. 300 x 250 um, stipe up to ca. 300 um high, non-
ostiolate, uniloculate, black, upper surface of dry asco-
Fig. 1. Lahmia kunzei. Two ascomata, one unopened and one
opened. Coll.: UME 29061. Magn.: x100.
349
mata with small fissures in SEM (Fig. 1).
Wall composed of a net-work of hyphae (textura intrica-
ta) with thick, scleroplectenchymatous walls, and inter-
cellulars filled with a gelatinous substance, strongly
melanized around surface hyphae (Fig. 2B), greyish green
under the surface layer and hyaline near the hamathecium.
Mature ascomata opening and exposing the asci by the
radial and irregularly splitting and partial disintegration
from centrum outwards of the upper part of the wall; empty,
Fig. 2. Lahmia kunzei. (A) Ascomata in a
fissure in aspen bark. (B) Ascomal wall in
surface view. (C) Ascoma, dry. (D) Ascoma
(detached), in water. (E) Ascomata (in
water), schem. ill., med. sect., upper one
showing branching of paraphysoids (asci
left out), lower one empty - Coll.: (A)
UPS: Liigenheim, 1861, leg. Rehm. (B, E) UME
29061. (C, D) UPS: Ibbenbiiren, on Populus
alba, leg. Lahm. - Magn.: (A) x8.5. (B)
<1400. (Cy) D) X30% (Bb) X10.
350
very thin-walled, cup-shaped ascomata (Fig. 1) with whitish
wall flaps at the rim, often seen with unopened and opened,
mature ascomata.
Hamathecium in young ascomata (before asci are to be
seen) consisting of filiform, anastomosing paraphysoids,
ca. 1-2 pm diam., directly continuing into the hyphae of
the surrounding tissues without any visible change in
anatomy. Paraphysoids streching between and above asci
during ontogenesis and here with much fewer anastomoses
than in basal part of hymenium; uppermost part of hama-
thecium secondarily producing paraphysis-like filaments
with free, slightly swollen tips, ca. 2-3 pm diam. (Fig.
3F), densely agglutinated by a thin covering layer of
faintly brownish, amorphous grains; hymenial gel around
paraphysoids in mature ascomata inconspicuous (stainable
with, e.g. Brilliant Chresyl Blue (BCB) in water mounts).
Asci clavate, very long-stalked, with hoof-like or
bifurcated bases, bitunicate, but probably non-fissituni-
cate, ectotunica thin, endotunica thin in asci with mature
spores, thick in broken asci (Fig. 3B), in upper part of
immature asci and in mature asci treated with KOH, with a
small ocular chamber, but without distinct ring structures,
(in mounts stained with BCB, a central cylindrical portion
stains less strongly than surrounding parts), 8-spored,
maturing almost synchronously, I-, 85-110 x 10-14 wpm,
easily breaking at the bases and squeezed out in water
mounts.
Ascospores hyaline, crescent-shaped, contorted, (1-)3(-
4)-septate, 18-37 x 4-5 pm, thin-walled, without distinct
perispore (Fig. 3 C,D,E).
DISCUSSION
Morphology
Lahmia kunzei is characterized by erumpent, stalked,
truncate, initially closed, hemiangiocarpous ascomata
breaking up irregularly by the splitting and partial
disintegration of the upper wall at maturity, ascomal wall
of textura intricata, hamathecium of paraphysoids with
secondary paraphysis-like ends, morphologically bitunicate
asci with very long and narrow stalks, easily breaking at
the base and becoming loose, and spirally twisted, finally
several-septate ascospores. Funk (1976: 868) gave a good
description of this species (as Parkerella populi), and
presented several excellent photographs. The Swedish
material agrees well with the protologue of Funk’s new
species, and there can be no doubt about their being
conspecific. There are a few small differences, however,
Fig. 3. Lahmia kunzei. (A). Ascus, not quite
mature (spores left out). (B). Broken ascus
with swollen endotunica. (C).Ascus with mature
spores. (D). Immature ascospores. (E). Mature
ascospores (F). Paraphysis-like tips of hama-
thecium. Coll.: (A) UME 29061. (B,D) UME 29063.
(C,E,F). UME 29068.
ool
552
which deserve some comments.
According to Funk (1l.c.) the ascomal wall should be
composed of textura epidermoidea. A more accurate descrip-
tion is textura intricata with cells separated by a gela-
tinous substance.
Funk (l.c.) described the paraphyses to be "not swollen
at the tips, 1-2 ym diam." Actually, they are a little
Swollen and ca. 2-3 um at the tips, whereas the rest of the
filaments in the hamathecium are only ca. 1-2 ym diam. This
may explain the discrepancy between Nannfeldt’s and Rehm’s
observations (see Nannfeldt 1932: 325). The hamathecium in
Lahmia kunzei is remarkable, and I have examined sections
and squash mounts of numerous immature and mature ascomata
in order to check whether there really are paraphyses with
free ends and not merely broken paraphysoids. There is no
doubt that there are. The ends are rounded and the cells
are often of uneven thickness. Young, immature ascomata
contain only paraphysoids, and the paraphyses must develop
secondarily from paraphysoids in the upper part of the
hamathecium during the ontogenesis. In mature ascomata the
Slightly swollen tips are glued together by faintly brown-
ish, amorphous, covering grains.
According to Funk (l.c.), the asci should be unituni-
cate, but with a markedly thickened ascus tip, giving the
appearance of a bitunicate ascus. "At dehiscence, the
contents of the ascus compress the thickening and burst
through the ascus wall, which shows some elasticity at this
point but which has no preformed pore". Nothing is men-
tioned about asci becoming loose and squeezed out from the
ascomata. It is possible that the whole ascus functions as
a diaspore, and that it opens in some way outside the
ascoma. Morphologically the ascus is bitunicate, but it may
have lost the fissitunicate function.
Systematical position
Problems immediately arise when attempting to refer
this species to an order in any of the recent classifica-
tions of the ascomycetes. Eriksson & Hawksworth (1985)
accepted 43 orders in their latest "Outline of the asco-
mycetes". Comparisons with Lahmia gives the following
results.
Orders 1-8 consist of discolichens, most of which have
disc-like apothecia, others with hysteriform lirellae, a
few with perithecioid apothecia. Nannfeldt (1932: 324)
stated that Lahmia was originally referred to the lichens,
but that Rehm (1890: 341) treated it among non-lichenized
fungi. Nannfeldt meant that the ascomata were built as
5538
those in Lecanorales, although they do not exhibit the
positive Iodine reaction. The ascomata of Lahmia kunzei are
hemiangiocarpous. This type of development characterizes
the orders 3-8, but in none of these lichen orders do the
ascoma, hamathecium (paraphysoids + "paraphyses"), asci,
etc. show any overall agreement with the structures in
Lahmia. Lahmia can not be a non-lichenized member of any of
these orders. Any similarities with true lichens are casual
only. Thus, superficially similar asci (though I+) with
spirally twisted, nematode-like spores occur in the genus
Sarrameana (Vezda & James 1973: 305), but the ascomata are
typical, gymnocarpous, disc-like apothecia in this genus.
Ascomata with stalked, globose fertil head occur in order 8
(Caliciales). In fact, Lahmia was originally referred to
this group of lichens by K6rber, but none of the genera
accepted by Tibell (1984) in Caliciales has the same type
of ascomal wall, hamathecium, asci, or spores as Lahmia.
Orders 9-14 contain non-lichenized discomycetes (the
Schaereriaceae, tentatively placed in Pezizales, and Baeo-
mycetaceae, in Leotiales, are lichenized). Order 9 (Peziz-
ales) differs, e.g in having usually large, fleshy, bright-
coloured apothecia, usually operculate asci, and hama-
thecium of paraphyses. In the Sarcosomataceae the para-
physes anastomose at the bases, but there are no other
Similarities with Lahmia. Some species in the order have
secondarily closed ascomata, but they are all large and
truffle-like and therefore inconsistent with Lahmia in this
context. Order 10 (Elaphomycetales) are truffle-like fungi.
Order 11 (Cyttariales) are parasites on Nothofagus, with
quite different ascomata, hamathecium, asci, etc. Order 12
(Leotiales) contains the bulk of the inoperculate, asco-
hymenial discomycetes. The ascomata are usually typical
apothecia with naked hymenium of asci and paraphyses, never
paraphysoids. Order 13 (Medeolariales) contains only one
species, a parasite on leaves of Medeola, lacking differ-
entiated ascomata, having true paraphyses, dark-coloured,
ornamented ascospores, etc. Gordon (1966, 1968) recognized
three types of centrum development in the Hypodermataceae,
i.e. the bulk of order 14 (Rhytismatales). In type I (Lo-
phodermium juniperinum (Fr.) De Not.) the hamathecium
consists of paraphysoids in young ascomata. In a later
stage they become free from the covering clypeus and
develop paraphyses from the free ends. This development is
thus similar to that in Lahmia, but there are so many
differences between these taxa that a close relationship is
very improbable. In Rhytismatales, the ascomata are im-
mersed in a stroma or host tissue and are often provided
354
with a clypeus, but they are never stalked as in Lahmia.
The mature hamathecium consists exclusively of paraphyses,
not mainly of paraphysoids as in Lahmia. The ascus morpho-
logy varies greatly within the order (Minter & Cannon 1984:
65), but never agrees with the morphology of the Lahmia
asci. In fact, Lahmia has no close resemblance to any of
the members in orders 9-14, and the genus can not be
included in any of these orders.
Orders 15-28 are unitunicate or secondarily pseudo-
prototunicate pyrenomycetes. Rather thick-walled, func-
tionally unitunicate asci occur in some representatives of
order 15 (Diaporthales), but the endotunica usually con-
tains an apical ring and the asci are not long-stalked as
in Lahmia. Moreover, paraphysoids and initially closed,
turbinate ascomata are unknown in that order. Order 16
(Sordariales) includes in current classifications the order
Coronophorales. Funk (1976: 871) referred his new genus
Parkerella (= Lahmia) to that order (= Nitschkiaceae in
Sordariales). He stated that it resembles Thaxteria in its
long-stalked asci with 3-septate spores. He pointed out
that the ascomata are turbinate in both genera, but the
tissues are pseudoparenchymatous in Thaxteria and not
plectenchymatous. Moreover, there should be paraphyses in
Parkerella and no distinct "Quellkérper" above the asci. As
mentioned above, the hamathecium consists of paraphysoids
and short secondary paraphysis-like filaments in Lahmia,
but this type of hamathecium does not occur in the Nitsch-
kiaceae. Moreover, in all members of the latter family the
ascomal wall consists of large, 2 isodiametric, dark-
coloured cells with characteristic "Munk pores" (inner
cells may be smaller and hyaline). The similarities in
ascus shape and spore septation are only casual, and Lahmia
is certainly not related to either Nitschkiaceae or any
other family in Sordariales. Orders 17 and 18 (Microascales
and Ophiostomatales) contain small perithecioid ascomata
without hamathecium, small evanescent asci, small non-
septate ascospores, etc. Order 19 (Diatrypales) are ostiol-
ate, commonly stromatic pyrenomycetes with I+ ring in the
asci, and non-septate, allantoid spores. Order 20 (Xylari-
ales) have ascohymenial, perithecioid ascomata, as a rule,
with I+ ring in the asci with dark-brown spores with germ
slit. Order 21 (Amphisphaeriales) are also ostiolate,
ascohymenial pyrenomycetes, differing from the previous
order in ascus and ascospore morphology. Most species in
order 22 (Halosphaeriales) are marine and have perithecioid
ascomata with usually evanescent hamathecium and asci. The
spores are often provided with characteristic gelatinous
395
equipments. Rather long-stalked asci occur in some Calo-
sphaeria species, order 23 (Calosphaeriales), but again,
the hamathecium is different (broad and tapering para-
physes), asci are attached to basal parts of paraphyses,
ascomata are perithecioid and have a periphysate ostiole,
etc. Order 24 (Trichosphaeriales) have ostiolate peri-
thecia, which are setose in many genera. The hamathecium is
lacking or may consist of paraphyses and periphyses.
Paraphysoids are reported for the genus Porosphaerellopsis
(Samuels & Miiller 1978: 128, as Porosphaeria), but there is
a periphysate ostiole, and the asci and spores are quite
different from those in Lahmia, and these taxa are very
remote. Order 25 (Phyllachorales) contains ostiolate,
ascohymenial, immersed pyrenomycetes. In order 26 (Hypocre-
ales) the ostiolate, light-coloured perithecia are usually
immersed in a stroma. Asci are thin-walled and _ short-
stalked, and the hamathecium usually consists of periphys-
oids, not paraphysoids. Order 27 (Eurotiales) have non-
ostiolate ascomata, but they are usually not stalked, and
the centrum is quite different from that in Lahmia, with
small globose, pseudoprototunicate asci, and many other
differences. Order 28 (Clavicipitales) are light-coloured,
usually stromatic fungi. The perithecia are ostiolate and
the very long and narrow asci have a semi-globose thicken-
ing in the top, pierced by a narrow pore. From the above it
is clear that Lahmia is very different from all pyreno-
mycetes with functionally unitunicate asci.
Orders 29-34 contain bitunicate ascomycetes. Lahmia
kunzei may be a bitunicate ascomycete, which has lost the
"Jack-in-the-Box" opening mechanism. Hamathecium of para-
physoids is known in all six orders, except for in Verru-
ecariales (periphysoids). However, the ascomata are apo-
thecioid in three of the orders (29-31), peritheciod in the
three others (32-34), but in none of them are they turbi-
nate, closed and open irregularly. Philippsiella atra Cooke
(Philippsiellaceae, Patellariales) has a superficial
resemblance, but the anatomy of the ascomata and the small
ellipsoidal asci are quite different. Seuratia spp. (Seura-
tiaceae, ?Arthoniales) have a thallus with cells embedded
in thick gel, but the genus is certainly not closely
related to Lahmia. Bundles of asci are laid down anywhere
in the thallus, and they are not enclosed in a hamathecium
and there is no ascomal wall separating the asci from the
vegetative thallus. Lahmia is out of place also in the
large and heterogeneous order Dothideales. Coccoidea Henn.,
parasitic on Quercus leaves, has circular, truncate asco-
mata, penetrating the host with a stipe (Eriksson 1981:
356
45), but the resemblance is superficial. The ascomata are
multi-loculate, open by pseudostioles and have pseudo-
parenchymatous walls. The asci resemble those of Lahmia a
little, but they are much more short-stalked, extrude a
part of the endotunica upon dehiscence, and contain ovoid
apiospores. Another family to consider is Tubeufiaceae. The
hamathecium of Tubeufia consists of paraphysoids (Samuels,
Rossman & Miller 1978: 184). The asci are stalked and
contain several-septate ascospores. However, the ascomata
open by a pseudostiole, and the surface cells of the
ascomal wall are circular, elliptic or angular in outline.
All species seem to have an anamorph.
Orders 35-43 (yeasts, Taphrinales, Laboulbeniales,
etc.) can all be left out of consideration. None of the
species in these orders has the slightest resemblance to
Lahmia kunzei.
The conclusion that follows from this discussion is
that Lahmia can not be included in any of the currently
recognized orders. The genus must be referred to a new
family in a new order. It was listed among genera of un-
certain affinities by Eriksson & Hawksworth (1985: 49-55).
None of the other genera in this large group seems to be
closely related to Lahmia. Therefore, a new monotypic
family and order is described.
I am not sure whether it is possible to use the names
Lahmiaceae and Lahmiales, as there are the older homonyms
Lamiaceae and Lamiales, based on Lamium L. (Phanerogamae).
The only other names which could be considered for basio-
nyms are Parkerella A. Funk, Parathalle Clem., and Lahmio-
myces Cif. & Tomas. As mentioned above, there is an older
homonym of Parkerella A. Funk. Parathalle is no longer
included in Lahmia. The type species, P. fueistingii
(Kérber) Clem. (= Lahmia fueistingii Kérber), is a synonym
of Arthrorhaphis grisea Th. Fr. (Poelt & Hafellner 1976:
219). Lahmiomyces, based on Lahmia picea Anzi, is not re-
lated to Lahmia kunzei (cf. Rehm 1890: 343). It is a member
of the Patellariaceae. The anatomy of the ascomal wall
resembles that in Lahmia, but the asci and the paraphysoids
are arranged in a hymenium covered by a thin epithecium.
The asci do not become loose in mature ascomata. The genus
will be treated more in detail in another paper.
This means that there is no generic name available for
constructing the names of the higher taxa if Lahmiaceae and
Lahmiales are considered homonyms of Lamiaceae and Lami-
ales. The ICBN gives no instructions about how to proceed
with homonyms of family names. I have decided to use the
names Lahmiaceae and Lahmiales, but this case should be
go”.
discussed by the General Committee of the Nomenclature
Section at the Berlin Congress in 1987, and if these names
are not accepted, I suggest that they are changed into
Mycolahmiaceae and Mycolahmiales (based on Lahmia prefixed
with "Myco-").
Lahmiaceae 0. Eriksson fam. nov.
Ascomata stipitata, turbinata, non ostiolata, uniloculata.
Paries e textura intricata compositus, fissuris radialibus
inaequalibus dehiscens. Hamathecium e paraphysoidibus et
apicem versus e filamentis paraphysium similibus compo-
situm. Asci bitunicati, longe stipitati. Sporae hyalinae,
septatae.
Typus: Lahmia Korber
Lahmiales 0. Eriksson ordo nov.
Characteres Lahmiacearum praebentes.
Typus: Lahmiaceae 0. Eriksson
Lahmiaceae 0. Eriksson fam. nov.
Ascomata stalked, turbinate, non-stiolate, uniloculate.
Wall of textura intricata, opening irregularly by radial
fissures. Hamathecium of paraphysoids and apically of
paraphysis-like filaments. Asci bitunicate, long-stalked.
Ascospores hyaline, septate.
Type: Lahmia Korber.
Lahmiales 0. Eriksson ordo nov.
Characters as in Lahmiaceae.
Type: Lahmiaceae 0. Eriksson
BIOLOGY
The Swedish specimens of Lahmia kunzei on Populus tremula
(closely related to P. tremuloides; see, e.g. Kriissman
1962: 227) were only found on rough bark in fissures on
aspen stems. Ascomata were never found on smooth bark or on
naked wood in fissures. The fungus was reported (as Parker-
ella populi) on cork bark of Populus tremuloides from the
central interior of British Columbia, Canada, and it was
usually found in association with Rhytidiella baranyayi
(Funk 1976: 868). The latter fungus was described by Funk
& Zalasky (1975: 752) and believed to be the real cause of
the cork bark disease. The etiology of the disease on P.
tremula deserves more studies. Lahmia kunzei is certainly
not the cause of the cork bark and the fissures. It is
358
wide-spread in Europe. I have seen herbarium material (UPS)
from Italy in the South to Finnmark in northern Norway in
the North. According to Rehm (1890: 342), the fungus occurs
on bark of Populus alba and P. tremula, but also, though
more rarely, on Salix, Robinia and Quercus. In the Nordic
countries it has only been found on Populus tremula. Two
samples of L. kunzei on P. alba collected by Lahm in
Germany are filed in UPS. There are no morphological
differences between ascomata from P. alba and P. tremula.
SWEDISH MATERIAL STUDIED
(1). Vasterbotten, Vaénnds par., ca. 1 km E. of Harrselsfors
power station, E. slope of the Ume River, 6.IX. 1985, leg.
O.E. (UME 29061); 18.V.1986 (UME 29068). - The fungus
occured on trees in a sheltered position at the base of a
steep slope. There were about 10 trees, all showing symp-
toms of cork bark disease. Some of the trees had very
extensive fissures in the bark, with cork bark near naked
wood. Several other fungi occurred on the bark, e.g. Tropo-—-
sporella fumosa Karsten. Aspens in a ravine near this
locality were all healthy.
(2). Vasterbotten, Umea, Sdérfors, mixed forest (Betula,
Populus, Picea, etc.) at the Ume River, 27.IV.1986, leg.
O.E. (UME 29063), 17.V.1986 (UME 29064). - Cork bark was
seen around dead twigs and large fissures on the stem. The
ascomata of Lahmia kunzei were not quite mature in April
1986, but contained mature, 3-septate spores in mid-May.
(3). Vasterbotten, Umea, Brd&dnnland, steep, sunexposed
slope at the Ume River near the outlet of Rupbdcken,
18.V.1986, leg. O.E. (UME 29065);
(4). Vadsterbotten, Umea, Between Ytterréda and Helene-
borg, 19.V.1986, leg. O.E. (UME 29066, 29067). In locality
(4) Lahmia kunzei was found on two aspens in a _ spruce
forest, very near the place where Rhytidiella moriformis
was encountered in 1975. The large aspen trees with this
fungus could no longer be found, and most of the spruces
have also been removed. Rhytidiella moriformis was searched
for, but could not be refound.
Numerous other collections of Lahmia kunzei from other
provinces in Sweden (S, UPS) were examined and verified
with the dissecting microscope.
ACKNOWLEDGEMENTS
I am grateful to Prof. Dr. R.P. Korf for the loan of North
American material of Parkerella populi, to Prof. Dr. R.
Santesson, Dr. L. Holm, and Dr. V. Shindler for review-
ays 08)
ing the manuscript, to Dr. L. Holm for correcting the Latin
diagnosis, and to Dr. Yue Jing-zhu for drawings (Fig. 2 A,
C, D).
LITERATURE CITED
ERIKSSON, 0. 1981. The families of bitunicate ascomycetes.
- Opera Bot. 60: 1-220.
- & HAWKSWORTH, D.L. 1985. Outline of the ascomycetes
- 1985. - Systema Ascomycetum 4: 1-79.
FARR, E.R., LEUSSINK, J.A. & STAFLEU, F.A. 1979. Index
Nominum Genericorum (Plantarum). II. - Regnum Veget.
1013763 1=127 6.
FUNK, A. 1976. Parkerella, a new genus of Coronophorales. -
Can. J. Bot. 54: 868-871.
- & ZALASKY, H. 1975. Rhytidiella baranyayi n.sp., associ-
ated with cork-bark of aspen. - Can. J. Bot. 53: 752-
159%
GORDON, C.C. 1966. Ascocarpic centrum ontogeny of species
of Hypodermataceae of conifers. - Amer. J. Bot. 53:3
319-327.
- 1968. Ascocarpic centrum ontogeny of species of Hypo-
dermataceae of conifers. II. - Amer. J. Bot. 55: 45-52.
KORBER, G.W. 1859-1865. Parerga Lichenologica. - Breslau.
KRUSSMANN, G. 1962. Handbuch der Laubgehélze. II. - Parey,
Berlin & Hamburg, 608 pp.
MINTER, D.W. & CANNON, P.F. 1984. Ascospore discharge in
some members of the Rhytismataceae. - Trans. Br. mycol.
Soc. 83: 65-92.
NANNFELDT, J.A. 1932. Studien tiber die Morphologie und
Systematik der nicht-lichenisierten, inoperculaten Dis-
comyceten. - Nova Acta R. Soc. Scient. upsal., ser. IV
8(2): 1-368.
POELT, J. & HAFELLNER, J. 1976. Die Flechte Neonorrlinia
trypetheliza und die Familie Arthrorhaphidaceae. -
Phyton 17: 213-220.
REHM, H. 1887-1896. Die Pilze. Ascomyceten: Hysteriaceen
und Discomyceten. - In: Rabenhorst, L., KryptogFlora
Deutschl., Osterr. und Schweiz 1(3): I-VIII, 1-1275.
SAMUELS, G.J. & MULLER, E. 1978. Life-history studies of
Brazilian ascomycetes 1. Two genera of the Sphaeriaceae
having, respectively Sporoschisma-like and Codinaea
anamorphs. - Sydowia 31: 126-136.
-, ROSSMAN, A. & MULLER, E. 1978. Life-history studies of
Brazilian ascomycetes. Three species of Tubeufia with,
respectively, dictyosporous/pycnidial and helicosporous
anamorphs. - Sydowia 31: 180-193.
360
TIBELL, L. 1984. A reappraisal of the taxonomy of Calici-
ales. - Beih. Nova Hedwigia 79: 597-713.
VEZDA, A. & JAMES, P. 1973. Sarrameana paradoxa A. Vezda &
P. James gen. nov. et sp. nova, eine bemerkenswerte
Flechte aus Neu-Kaledonien. - Preslia (Praha) 45:
305-310.
ZALASKY, H. 1968. Rhytidiella moriformis n. gen., n. sp.
causing rough-bark of Populus balsamifera. - Can. J.
Bot. 46: 1383-1387.
- 1975. Cell deformities in bark and sapwood caused by
Rhytidiella moriformis and Keissleriella emergens
infections in poplar. - Can. J. Bot. 53: 780-783.
MYCOTAXON
VOLE AXVEL pp. S61-375 October-December 1986
CULTURAL STUDIES IN TUBULICRINIS AND XENASMATELLA
(CORTICIACEAE, BASIDIOMYCETES )
Nils Hallenberg
Department of Systematic Botany
Gothenburg University, S-413 19 Géteborg, Sweden
ABSTRACT. Cultural characters are reported for six species
CGuimouLieninis ) Cl. borealis, .T.. | calothrix, T. .gracil-
limus, T. medius, T. strangulatus, T. subulatus) and
two species of Xenasmatella (X. allantospora, X. tulas-
nelloidea). MIntercompatibility tests between a number
of specimens have been carried out to assess the variation
in substrate and habitat preference and geographical dist-
ribution. Interincompatible sibling species have been found
in five of the studied species.
INTRODUCTION. Very little information exists on cultural
characters and intercompatibility matings in the genera
Tubulicrinis and Xenasmatella (Boidin 1958, Hallenberg
1984). Consequently, such data has hitherto been unavailab-
le for taxonomical decisions.
The present investigation reemphazises the homogenity with-
in respective genera as far as the investigated species are
concerned. Further, the results from intercompatibility
tests have been used to assess variation in fruitbody mor-
phology, substrate and habitat preference and geographical
distribution. Several sibling species were detected within
the investigated material, but no new species are describ-
ed. In this study, groups of compatible specimens are con-
sidered sibling species when they are genetically isolated
from each other (interincompatible) and when there are no
significant morphological differences between their fruit-
bodies. All siblings found here will be determined as res-
pective "morphological species" when using the keys’ const-
ructed. by, Eriksson et.al. (1987) and Jiilich (1984).
It is important to obtain more knowledge of sibling species
complexes for the understanding of speciation processes.
Before describing the entities as species with Latin bino-
mials it is necessary to have a more precise knowledge of
characters other than morphological ones (substrate pre-
ference, geographical distribution, etc.). Moreover, I do
not see any contradiction in accepting a "biological spe-
cies concept" and application of a "practical standard of a
species" as defined by Parmasto (1985). The latter term
362
reflects, the) present knowledge of) a:species. an thic, papers
the different siblings will be referred to different "ty-
pes" within a (morphological) species.
This investigation is'made in: * connection "with — thes f.ora
project "Corticiaceae of North Europe" and references con-
cerning the treated species are given in this flora (Eriks-
sOnuete al =. cLIOGIO
METHODS. The specimens originate in Austria, Canada, Den-
mark, Norway, Romania, and Sweden. They have been carefully
compared with types or other representative material. When
spore measurements are given these have been made from spo-
re prints. Single-spore (SS)-cultures and polyspore (PS)-
cultures were isolated after dispersal on common malt agar.
To determine the breeding ability (i.e., the ability to
form clamps “in matings) ‘of the individidal SS=cultures,
matings between SS-cultures of each specimen were first
made. Compatible mating types from one specimen were mated
with compatible mating types from the other specimens. In
some cultures it was impossible to obtain compatible mating
types. However, such SS-cultures have been accepted if they
were compatible in matings with cultures from other speci-
mens. Further, in other specimens it was possible to indi-
cate the polarity pattern. Formation of clamps in matings
was looked for after that the mycelia had been in contact
for 6-8 weeks.
Culture characters of the PS-mycelia were analyzed after 3
and 6 weeks’ growth. Culture codes are from Nobles (1965)
with emendations by Boidin and Lanquetin (1983). Drop tests
for extracellular oxidases (Syringaldazine, Gum Guaiac,
Guaiacol, 1-Naphtol, p-Cresol, L-Tyrosine) follow Marr
(1979). Nuclear staining was made with Giemsa according to
Boidin (1958). Terminology of nuclear behaviour refers to
Boidin and Lanquetin (1984) as well as earlier reports by
Boigin.
In the lists of "material" (below, under each species) in-
formation concerning the studied specimens are given. Such
data are cathegorized by slanted lines: Culture number/
substrate/ locality/ number of available SS-(PS-)cultures
or SS-numbers of compatible mating types. Compatible mating
types are separated by a Slant’ line’, "/".
Cultures’ are” ‘stored in’ the” culture-—collection\= atvpene
Department of Systematic Botany, Gothenburg University. The
GB-numbers refer to the culture-collection. The original
specimen together with spore-print can be identified by the
same number and are kept in the herbarium (GB).
363
RESULTS OF “CULTURAL STUDIES ANDY GCOMPATIBIGITY TESTS.
Tubulicrinis borealis John Erikss.
Material:
GB 339/ Picea/ Sweden, Narke/ 4 SS.
GB 439/ Picea/ Sweden, Vastergdtland/ 1/4.
GB 795/ Pseudotsuga/ Canada, B.C., Vancouver Island/3 SS.
Compatibility between these specimens has previously been
shown (Hallenberg, 1984). In the present study, these cul-
tures proved incompatible with specimens representing the
chosely “related “species J.) strangulatus. Altogether 26
negative confrontations recorded.
According to Eriksson et al. GOS fice a be MDOT eC aek tis Nel aS so
mainly northern distribution in North Europe and occurs on
decorticated, coniferous wood (as did the tested speci-
mens). It seems to be associated with brown-rotted wood.
CULTURE CHARACTERS (GB 439): Fags. AGIA
Aerial mycelium absent to downy (in old parts). Advancing
zone even, appressed. Ordinary branching hyphae, clamped at
ais septa, with "thin to slightly thickened walls, 1.5=3 ~ym
wide in the margin , up to 7 pm wide in old parts and part-
ivewieny irreéeular outlines (rig. 2A) no staining in), Mel-—
Zeer.
Colony radius 21 mm after 6 weeks.
CODE:
ee Crea LON). Yous SON esos 44. eo 410d s
OXIDASE REACTIONS: All tests negative.
POLARITY: Heterothallic, type of polarity unknown.
CYTOLOGY: SS-mycelium with uninucleate cells, PS-mycelium
GECaALYOULEC ;
NUCLEAR BEHAVIOUR: Normal.
T. calothrix (Pat.)Donk
Material:
GB 341/ Pinus/ Sweden, Smaland/ 1,3,4/2,5.
GB 357/ Populus/ Sweden, Uppland/ 1,2,3/6,7.
Compatibility has been found between these two specimens.
Morphologically they are in accordance with the descrip-
elons civenopy Erikssonmet «al. 7'(1987).. Le *\ calothrix. as
there reported as fairly common in N. Sweden, becoming more
rarewtowards: the (SS. "parts.
The specimens in this compatibility group were collected in
thesc. iparts.or No Europe. Lt ts interesting to “note ‘that
GB 357 was collected on a substrate other than coniferous
wood.
CULTURE CHARACTERS (GB 357): Fae, (Girt
Aerial mycelium absent. Advancing zone even, appressed.
Ordinary branching hyphae, with clamps at all septa, with
364
thin to slightly thickened walls, 1.5-6 wm wide, with scat-
tered capitate hyphal endings in old mat (globule up to 13
pm in diam.). Moreover, hyphae in the old mat are frequent-
ly somewhat moniliform (fig. 2B). At some septa, constric-
tion /of (the) hypha, (fig. 2B) is) found, which! ts. 4 ifeatune
commonly found in Hyphodontia. No reaction with Melzer.
Colony radius 18 mm after 6 weeks.
CODE:
28) (SCE UL C2 igi 2isc MSOs) Os.) 4), eo dO eE yO Olen noi.
OXIDASE REACTIONS: Syringaldazine, 1-Naphtol, Gum Guaiac,
and Guaiacol positive. p-Cresol and L-Tyrosine negative.
POLARITY: Heterothallic, type of polarity unknown.
CYTOLOGY: SS-mycelia with uninucleate cells, PS-mycelium
dicaryotze,
NUCLEAR BEHAVIOUR: Normal.
Lu graciilinus. (Roe. .& Jacks,)G.H. Cunn.
Syn.: T. glebulosus (Bres.)Donk
Material:
GB 417/ Pinus/ Austria, Steiermark/ 1 PS.
GB 285/ Betula/ Sweden, Torne Lappmark/ 1,4,5/2,6.
GB 787/ Populus/ Canada, Quebec/ 2/3.
GB 912/ Betula/ Sweden, Lycksele Lappmark/ 1,9,10/3.
GB 1251/ Picea/ Norway, Oppland/ 4/5.
GB 1253/ Picea/ Norway, Oppland/ 3/2,4,5.
Three compatibility groups have been discerned between
which there is complete incompatibility: A) GB 417; B) GB
1:2'5.1,,),GB) 1253%)-C) GB.285..°GB 787... .GB 912s.) Altogether,
negative confrontations recorded.
Only minor morphological differences exist between these
three groups and in the keys of Eriksson et al. (1987) and
Julich (1984) all will be determined as). T...gractlisanus,
Although very little material is available the following
separating characters of the fruitbodies may be useful in
the search for delimitation of the sibling species:
A) Substrate: coniferous wood (Pinus). Spores size: 8,5-9.5
X 2 um. Subiculum and context is mainly composed of thick-
walled skeletals and clamped generative hyphae with thicke-
ned walls, interwoven in a rather loose texture. There are
numerous transitions between these two kinds of hyphae as
well as between skeletals and lyocystidia. A high propor-
tion of the lyocystidia are enclosed, while some project
beyond the hymenial surface. Lyocystidia are moderately
amyloid, skeletals are weakly to not-amyloid, 3-4 um wide,
with a narrow lumen, occasionally branched.
B) Substrate:, coniferous:wood (Picea). Spore | size?) vets
1.8-2 pum. Fruitbodies rather thin; hyphae in subiculum and
context rather thin-walled, interwoven; skeletals absent.
Hyphae in the. subiculum and) adjacent part of \the context
may be weakly or not amyloid. Lyocystidia moderately amy-
loid:, distinctly, projecting.
365
C) Substrate: deciduous wood (Betula, Populus). Spore size:
6-8 x 1.8-2 pm. In the subiculum and adjacent part of the
context, scattered skeletals are found together with gene-
rative hyphae with thickened walls. These may all be weakly
or not amyloid. Context hyphae with a pronounced vertical
orientation. Lyocystidia rather weakly amyloid, embedded or
projecting.
The morphological differences, differences in substrate
preference, as well as the results from the intercompatibi-
lity tests indicate that these three groups represent dif-
ferent and distinct species. Hjortstam (1979) also reports
Wiae wNOnds comaberial wot) Tiieractilinus) jis heterogenous
and probably includes more than one taxon.
The sibling species reported here should be referred to as
"coniferous type A" (GB 417), "coniferous type B" (GB 1251,
GB 1253), and "broad-leaved type" (GB 285, GB 787, GB 912).
CULTURE CHARACTERS (GB 417 = coniferous type A):
Page Dea
Aerial mycelium downy, whitish granules in agar. Advancing
zone even, appressed. Ordinary branching hyphae, clamped at
alieicepita, wath thin, te slightly, thickened) walls) 125-7) orm
wide but a distinct difference between "narrow" and "wide"
ones. Thick-walled, ellipsoidal chlamydospores, 9-12 X 6-10
pm, with light refracting contents (with phase contrast),
foumed= in. chains, »;abundant ini the granules: (figt 26). No
reaction in Melzer.
Plates covered in 6 weeks.
CULTURE CHARACTERS ‘(GB 1253: = coniferous) type B; GB 912. =
broad-leaved type): Pai et ulik ys bus mee)
Aerial mycelium absent to downy (in old parts). Advancing
zone even to slightly bayed, appressed. Ordinary branching
hypuec. with clamps jatwallisepta, with thin) to “slightly
thickened walls, 1.5-4 um wide, constricted at some septa
(4cil4kexan Hy phodontia), in old. parts, some, hyphae .jirre-
gularly swollen) )Cup ito 11 pm wides fig. 2D). No reaction
with Melzer.
Cotonyi radius, after 6 weeks: (1.5. mm Cin. GB 912), 10° ‘mm. Gin
GBat 253).
CODES:
Be Weve i io eros) Se eh) So GMO ere MO Oe IO FES
SBue ce a MG De Oye tet N O00 56 5k Kilo tah Oil Oldie
GRO tee ae 2ID Nis. 2 OF nM opis MOO). ATK Oe OS's.) Ollie
Fig. 1. Cultures after six weeks growth. a) Tubulicrinis borealis, GB
439, b) T. medius, GB 976, c) dito, GB 338, d) T. gracillimus "con-
iferous type AY, GB 417, e) T. gracillimus "coniferous type B", GB
1253, f£) T. gracillimus “broad-Ieaved type", GB 912, g) T. calo-
thrix, GB 357, h) T. subulatus "mixed forest type", GB 1292, i) Xe-
nasmatella_allantospora "type A", GB 1315, j) Tubulicrinis strangula-
tus "moderately decayed type", GB 1056, k) T. subulatus "old spruce
forest type’, GB 1255, 1) Xenasmatella tulasnelloidea "Nordic type".
- Photos Ellen Hansson.
366
368
OXIDASE REACTIONS:
Syringald. 1-Napht. Gum G. Guaiacol p-Cresol
L-Tyr.
GB 417: s = S e rR =
GB 1253: + + + + (+) (+)
GB 952: + + + + (+) (+)
POLARITY: Unknown for GB 417 (no SS-cultures available);
heterothallic but type of polarity unknown in GB 1253 and
GB 912.
CYTOLOGY: SS-mycelia with uninucleate cells (GB 1253, GB
912), PS-mycelia dicaryotic.
NUCLEAR BEHAVIOUR: Normal (GB 1253, GB 912).
T. medius (Bourd. & Galz.)Oberw.
Material:
GB 338/ Picea/ Sweden, Smaland/ 1 PS.
GB 976/ Picea/ Norway, Hedmark/ 2,6/3,7,8.
Compatibility has been found between the two specimens.
According to Eriksson’) et, al. (1987), Dyocystidia winger,
medius may be variably amyloid, even though staining in
Melzer in most cases is strong. In the specimens investiga-
ted here, the lyocystidia are only weakly amyloid with the
exception|)\of ‘the part \ of the wall that is close ta the
lumen. This part is characteristically more strongly amy-
roid.
In other respects, the characters of these two specimens
coincide with the concept given in the flora.
CULTURE CHARACTERS (GB 338, GB 976): Figiavl, By Comoe.
Aerial mycelium downy, in old mats slightly cottony. Advan-
cing zone even, appressed. Ordinary branching hyphae, with
clamps at all septa, 1.5-4 wm wide; marginal hyphae thin-
walled, their apical parts frequently flexuous and with an
irregular outline (fig. 2F); hyphae in the old parts with
thickened walls, straight to somewhat flexuous, frequently
irregularly widened (up to 11 wm wide; fig. 2E), some hyph-
al segments in the old parts rather thick-walled and dis-
tinctly amyloid.
Colony radius 12 mm after 6 weeks.
CODE:
2a e) C2D Va eee COOLS Ve 26 ain Soe CAO eee « \ Arie hoy an bee Oren
OXIDASE REACTIONS:
Syringald. 1-Napht. Gum G. Guaiacol p-Cresol L-Tyr.
GB 338: (+) - + pe 4 es
GB 976: - - + - - -
POLARITY: Heterothallic but type of polarity unknown.
CYTOLOGY: SS-mycelia with uninucleate cells, PS-mycelia
dicaryotic.
NUCLEAR BEHAVIOUR: Normal.
369
2—— Pe a
Fig. 2. Culture details in Tubulicrinis (further explanations in
texey.) AVM Loe borealis, CB.) 4394 Bit. calothrix), CB. 35/7, C) D. gra-
cillimus "coniferous type A", GB 417, D) T. gracillimus "broad-
leaved type”, GB 912, E) T. medius, GB 338, F) dito, GB 976, G) im
subulatus "old spruce forest type", GB 1255, H) T. subulatus "mixed
forest type", GB 1292.
CYA)
Te strangulatuws:.Larss. <¢ Hjortet.
Material:
GB 905/ Picea/ Sweden, Lycksele Lappmark/ 1,4,5/2,6.
GB 978/ Picea/ Norway, Hedmark/ 1,3,5,6/8.
GB 1056/ Picea/ Romania, Suceava/ 2,3,5/6.
GB 1419/ Picea/ Norway, Oslo/ 8,9/5,6.
GB 1544/ Abies/ Romania, Neamt/ 1/5,6; 2,7,8/3,4.
Compatibility has been found between GB 978, GB 1056, GB
14}9% Sands CB 41544. WAL tthese ‘cultures were sincompau woe
with GB 905. Altogether 25 negative confrontations . recor—
ded.
The specimens agree in their morphology with the the type
specimen. and “the description (given “in y Eriksson, et ac.
(1987 \c,- Lieriley is known, <about. “the “distribution orem
strangulatus;) ~but “as 7indicated):bys the intercompatepisasr,
tests, the species is widespread and occurs in coniferous
forests,
int tthe sanorpholeey 7" ls istrangulatus “dippers from sly
borealis mainly by having slightly broader spores (5-5.5 ey,
2.5 pm versus 5.5-6 X 2-2.3 pm, when measured from’ spore-
prints) and an inamyloid basidial layer.
Further, there are no noticeable morphological differences
in the fruitbodies between GB 905 and the other specimens
treated here. On the other hand, there seems to be a diffe-
rence in substrate preference. While the specimens in the
main intercompatibility group have been found on moderately
decayed wood (brown-rot), GB 905 was found on_ strongly
brown-rotted wood. The decay with which T. strangulatus
is) associated istobviously. caused) by “Fomitopsise spimicetia
(Ere) Karst.
The sibling species ini TT. strangulatus should be referred
to as “moderately decayed type’ CGB 978, GB 1056,°CB 1419,
GB 1544) and “strongly decayed type" (GB 905).
CULTURE CHARACTERS) CGB: 1056): Pion kK
These agree with those for T. borealis (GB 439; see abo-
ve) with exception of slower growth in the former (14 mm in
6 weeks versus 21 mm). Moreover, inithe tests for extracel-—
lular oxidases, jno (reaction was seen in, Tes iborealic,
while some were tests were positive in T. strangulatus.
GODER 2a4 Bia vee C26) 8249-36. 84pm sarod
OXIDASE REACTIONS: Syringaldazine, 1-Naphtol, Gum Guaiac,
and Guaiacol positive. p-Cresol and L-Tyrosine negative.
POLARITY: Heterothallic, tetrapolarity indicated’ (GB 1544 )<
CYTOLOGY: SS-mycelia with uninucleate cells, PS-mycelium
dicakr yiotate:.
NUCLEAR BEHAVIOUR: Normal.
571
T. subulatus (Bourd. & Galz.)Donk
Material:
GB 288/ Picea/ Denmark, Jylland/ 1,3,6,7/2,4,5.
GB 363/ Populus/ Sweden, Uppland/ 3/5.
GB 914/ Pinus/ Denmark, Jylland/ 1,4/2.
GB 1073/ Pinus/ Scotland, Perthshire/ 5 SS.
GB 1292/ Picea/ Sweden, Halland/ 1/5,6; 3/2,4.
GB 1353/ Picea/ Sweden, Gotland/ 1/2.
GB 496/ Abies/ Canada, B.C./ 9 SS.
GB 1255/ Picea/ Norway, Oppland/ 1/4.
GB 1546/ Picea/ Romania, Brasov/ 1,2,4,8/6.
Two compatibility groups have been found:
PCD we oo, Gb JOG. 'Go, 914. GBT O7G.. Gb 1292. GB 1353's
DIEGO, GB 255), (GB 1546.
Complete incompatibility found in matings between represen-
tatives from each group (sibling species). Altogether 28
negative confrontations recorded.
According “to -the idelimitation of ‘I. subulatus .civen’” an
Eriksson et al. (1987), lyocystidia in the fruitbodies are
variable. Apices of lyocystidia may be subulate to mucrona-
te and their walls may be inamyloid - light greyish - weak-
ly amyloid, when treated with Melzer. This variation exists
in both of the compatibility groups treated here. The only
indications of differences between the siblings seem to be
in their habitat preference. While group I has been found
in mixed or cultivated forests, the two representatives of
group II were collected in old spruce forests in montane
regions’.
Group I should be referred to as "mixed forest type" and
group as “old spruce forest, type”.
CULTURE CHARACTERS (GB 1292 = "mixed forest type"):
Pow lise
Aerial mycelium downy. Advancing zone slightly fringed -
bayed, appressed. Ordinary branching hyphae, with clamps at
addi septa, thin-walled, 125-3./5 ‘um wide;cin “old “parts " of
the mycelium with frequent swellings on the hyphae, inter-
cabvanby -or terminally, wp to 18 um wide’.
Colony radius 5 mm after 6 weeks.
CULTURE: CHARACTERS (GB 1255°25"old spruce forest’ type"):
|S ae 2 ie ne
Aerial mycelium downy. Advancing zone even, appressed.
Ordinary “branching hyphee; with clamps2 at all ‘septa,
oes) mn wide, with thinv itor ‘slightly thickened walls;
hyphae in the margin uniform in diameter, in old parts
Somewnat. irregular;: lyocystidia found in: ‘old ‘parts of; ~ the
mycelium, more or less capitate, non-amyloid.
Colony radius 12 mm after 6 weeks.
CODE:
Dee O22 ae 2D ee. Cnt? Ole D2) 9 be On Cr sy or irsh gis LOO carol
Ose Ds OO Sits a SO SD Oae a is OO «Os, ON.
one
OXIDASE REACTIONS: Both GB 1292 and GB 1255 were positive
in all) tests.
POLARITY: Tetrapolarity indicated for "mixed forest type",
heterothallism (polarity unknown) in "old spruce forest
by Dena.
CYTOLOGY: In both siblings, SS-mycelia have uninucleate
cells, PS-mycelia dicaryotic.
NUCLEAR BEHAVIOUR: Normal.
Xenasmatella allantospora Oberw.
Material:
GB 254/ Pinus/ Denmark, Jylland/ 1/2.
GB 372/ Picea/ Austria, Steiermark/ 3 SS.
GB 399/ Picea/ Austria, Steiermark/ 3/5.
GB 442/ deciduous wood/ Sweden, Vastergétland/ 6/9.
GB 1315/ deciduous wood/ Sweden, Gotland/ 1,6/7; 4/3.
Compatibility was demonstrated between GB 372, GB 399, GB
442, and GB 1315. All these specimens were incompatible
with GB 254 (16 negative confrontations recorded).
No noticeable morphological differences were observed bet-
ween the fruitbodies. Spore sizes (when measured from spore
prints) varied minutely between the specimens but all were
within the limits 4-6 X 1.5-2 pm.
All specimens were found on decayed, decorticated wood
either of coniferous or deciduous trees. The habitats were
coniferous forests except for GB 1315 (Fraxinus-Betula-
Quercus forest) and GB 254 (mixed forest on the sea-shore).
The major compatibility group should be referred to as "ty-
pe A" while GB 254 is "type B".
CULTURE ‘CHARACTERS. .CGBi 372... 1915) Fags) sist i Se
Aerial mycelium slightly cottony - subfelty. Advancing zone
a little bayed and fringed, appressed. Hyphae thin-walled,
sparsely branched in the margin, frequently simple septate,
0.8-1.5 pm wide; hyphae in old parts of the mycelium with
clamps at all septa, the clamps are often characteristical-
ly flattened, hyphae up to 2 pm wide, frequently densely
ramified, with scattered swellings. Brown resinous residues
in agar (old parts of mycelium).
Colony radius 8 mm after 6 weeks.
CODES) a) G2 ose ol lie 26.0982" 2e86), AB a at aoe One
Oi
OXIDASE REACTIONS: Syringaldazine, 1-Naphtol, Gum Guaiac,
Guaiacol positive. p-Cresol and L-Tyrosine negative, except
for GB 372 (L-Tyrosine positive).
POLARITY: Heterothallic, tetrapolarity indicated (GB 1315).
CYTOLOGY: SS-mycelia with uninucleate cells, PS-mycelium
regularly dicaryotic.
NUCLEAR BEHAVIOUR: Normal.
X. tulasnelloidea (Hohn. & Litsch.)Oberw.
Material:
GB 944/ Fagus/ Denmark, Jylland/ 5 SS.
GB 1164/ deciduous wood/ Sweden, Bohuslan/ 5 SS.
GB 1330/ deciduous wood/ Sweden, Gotland/ 1,2,5/6.
GB 1338/ Fraxinus/ Sweden, Vastergétland/1,3/2,5,6.
GB 1360/ Fagus/ Sweden, Dalsland/ 6 SS.
GB 406/ Abies/ Austria, Steiermark/ 1,4/2,3,5.
GB 1532/ Fraxinus/ Romania, Iasi/ 8 SS.
Celso /tedec, wood/) (Romania. ) Brasov/* \1,6/2;3)4,537.
Two compatibilty groups have been found:
teeter Se (GBT oO4h Gh 1330 Gbil358, GB 1360:
PEGA. 6Gh: LSs2) 77Gb bo3O.,
Complete incompatibility was found between these groups
(altogether 48 negative confrontations recorded).
No noticeable morphological differences were detected bet-
ween the fruitbodies. Spore sizes varied minutely between
the specimens of both groups but all were within the limits
4.5-6 X 3.5-4.3 um. All specimens were found on decayed,
decorticated, deciduous wood except GB 406, which was found
on the bark of Abies. Their habitats were fertile deciduous
forests, Fagus-forests or mixed forests with Fagus.
Specimens belonging to group I should be referred to as
"Nordic type", group II as "Central European type"
ee v
Fig. 3. Culture details in Xenasmatella (further explanations in
text). A) X. allantospora "type A", GB 1315. B - D, X. tulasnelloi-
dea "Nordic type", GB 1330; B) Facaneeeiy branched hypha, C) ampullate
clamps in old mycelium, D) chlamydospores.
374
CULTURE CHARACTERS) (GB +8330, (GB.21360 =e Nordtesty peje
Fig, IM .3B=)
Aerial mycelium absent towards the margin, farinaceous in
old parts. Advancing zone slightly bayed, appressed. Hyphae
clamped at all septa, thin-walled, clamps often characte-
ristically flattened )(f129.°63B); hyphae inthe maroin wer,
narrow, O.7-2> pm’ wide, racemose (fig: 3B)s) imsijold’ (parte
hyphae frequently widened, especially close to the septa
(fig. 3C). Conidia (chlamydospores) subglobose - broadly
ellipsoidal, 3-4 X'- 4-7. pm,.)uni-!’ to%bDinucheate?) rather
thick-walled and with light refracting contents (with phase
contrast), formed terminally on short side-branches (fig.
3D). Basidia and basidiospores most frequent in a zone bet-
ween the margin and the chlamysdospore-producing, farina-
ceous part.
Colony radius 20 mm after 6 weeks.
The cultural characters of xX. tulasnelloidea:) were also
studied “by Boudin GlLOssperwi th sinilanr vesuiits.
CODE:
ZAWMOCGHP tls. G42 POON ts S ee chilis NGOun Dandie Otros.
OXIDASE REACTIONS: Syringaldazine, 1-Naphtol, Gum Guaiac,
Guaiacol positive. p-Cresol and L-Tyrosin negative.
CYTOLOGY: SS-mycelia with uninucleate cells, PS-mycelium
dicaryotic, conidia binucleate.
NUCLEAR BEHAVIOUR: Normal.
CULTURE CHARACTERS (GB 1532 = "Central European type");
The only differences between this culture and those repre-
senting the "Nordic type" seem to be bigger chlamydospores
(7.5-12 x 4-4.5 pm versus 4.5-6 x 3.5-4.3 pm) and a_=e some-
what more rapid growth (35 mm versus 20 mm). No differences
were noted in oxidase reactions and cytology.
DISCUSSION
The cultural characters of the species studied to date
reaffirms the homogeneity within these two genera. The spe-
cies of Tubulicrinis have slow growth, hyphae with dis-
tinct ‘cell’ “walls... with clamps*at all septa) and) a norpmas
nuclear behaviour. These characters, as well as the featu-
res: of hyphae,” *make a relationship to Hyphodontia plau=
sible. This relationship is further confirmed by the fruit-
body morphology in the two genera.
Xenasmatella allantospora and X. tulasnelloidea were
been placed in different genera by Jiilich (1979). However,
it is obvious, fromithe culture © characters, that? sihey (ace
closely related: slow growth, very narrow hyphae which may
be racemosely branched, and characteristically flattened
clamps.
ACKNOWLEDGEMENTS
I am indebted to Ellen Hansson, G6teborg, who made the
laboratory work and the photos; to K. Seifert, Ottawa, who
corrected the English. Financial support was given by the
Swedish Natural Science Research Council.
othe
REFERENCES
Boidin, J. 1958: Essai biotaxonomique sur les Hydnes eae et les
Corticiés. Rev. Mycol. Mem. hors. Ser. 6, 388 p
Boidin, J., Lanquetin P. 1983: Basidiomycétes Nel TSU PAT ae épithe-
loides étalés. Mycotaxon 16 (2): 461-499.
Boidin, J., Lanquetin, P. 1984 a: Repertoire des donnees utiles pour
effectuer les tests d'intercompatibilite chez les Basidiomy-
cétes. I. Introduction. Cryptog. Mycol. Tome 5: 33-45.
Boidin, J., Lanquetin, P. 1984 b: Repertoire des donnees utiles pour
effectuer les tests d' intercompatibilité chez les Basidiomy-
cétes. III. Aphyllophorales non porées. Cryptog. Mycol. Tome
5: 193-245.
Eriksson, J., Hjortstam, Larsson, K-H., Ryvarden, L. 1987: The Cor-
ticiaceae of North Europe. Vol. 8 (in manuscript). Fungiflora.
Hallenberg, N. 1984: Compatibility between Species of Corticiaceae
(Basidiomycetes) from Europe and North America. Mycotaxon 21:
335-388.
Jiilich, W. 1984: Die Nichtblatterpilze, Gallertpilze und Bauchpilze.
Kl. Kryptogamenfl. IIb/1 Basidiomyceten. G. Fischer Verlag.
Marr, C.D. 1979: Laccase and Tyrosinase Oxidation of Spot Test Rea-
gents. Mycotaxon 9 (1): 244-276.
Nobles, M.K. 1965: Identification of Cultures of Wood-inhabiting
Hymenomycetes. Can. J. Bot. 43: 1097-1139.
Parmasto, E. 1985: The Species Concept in Hymenochaetaceae (Fungi,
Hymenomycetes). Proc. Indian Acad. Sci. (Plant Sci.), Vol.
94( 2-3): 369-380.
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MYCOTAXON
VODeONNV LL Spe otic 403 October-December 1986
FOLIICOLOUS ASCOMYCETES 7. PHYLOGENETIC SYSTEMATICS
OF THE CAPNODIACEAE
DON R. REYNOLDS
Natural History Museum
900 Exposition Boulevard
Los Angeles, California 90007 USA
ABSTRACT
The Capnodiaceae sensu lato is tested as a monophyletic
group with compatible character assisted parsimony
analyses. Nineteen ascocarpic and hyphal characters are
analyzed. The resultant phylogenetic inferences support
the recognition of the family Capnodiaceae (Saccardo)
Hohnel and three proposed subfamilies
Antennularielloideae, Capnodioideae, and Limacinioideae.
A. INTRODUCTION.
Capnodiaceous fungi are a group of ascomycetes which have
evolved as inhabitants of the aerial surface of living
plants. Both teleomorphic and anamorphic characters have
been used to characterize the species since the
Capnodiaceae was recognized. The Capnodiaceae sensu lato
has been broken into a number of smaller units, largely on
the basis of assumed modes of asexual reproduction.
The Capnodiaceae sensu lato was reviewed by Batista and
Ciferri (1963) in their monograph on the "Capnodiales".
The limitations of the family were based on the
antecedents of Arnaud (1911), Berkeley and Desmazieres
(1849), Ellis and Everhart (1892), Hohnel (1909, 1910),
Montagne (1849), Saccardo (1882), Spegazzini (1918), and
Theissen and Sydow (1917), Tulasne and Tulasne (1863).
Recent affirmations of this approach have been made by Arx
and Muller (1975), Luttrell (1973), and Reynolds (1971,
191 Og to) 9 A982). AIS oa).
The genera of the Capnodiaceae sensu lato were regrouped
by Hughes (1976) in an effort to carry out an avowed
predilection for small families (Hughes, 1972). Anamorphic
states were proposed as teleomorphic components of groups
of capnodiaceous genera on prima facie evidence. Certain
elements of the ascocarp centrum were reinterpreted in
order to affect a stronger affinity with non-capnodiaceous
taxa (Corlett 1970; Hughes, 1976). These groups were given
the family names Antennulariellaceae, Capnodiaceae,
Euantennariaceae, Metacapnodiaceae, and Triposporiop-
sidaceae. This concept has been endorsed by Barr (1979,
1983) who spread the taxa of the Capnodiaceae sensu lato
through two subclasses of the Loculoascomycetes. Eriksson
(1981, 1982) accepted this concept with reservation and
united these taxa into one suborder.
378
I have revised several capnodiaceous genera with a
teleocentric focus, i. e. with emphasis on the ascocarpic
characters. Several of the smaller taxa have been
restored to the Capnodiaceae as a result. For example,
Triposporiopsis spinigera (Héhnel) Hohnel, the type
species of its genus and the family Triposporiopsidaceae
Hughes (Hughes, 1976), was synonymized with Trichomerium
Spegazzini (Reynolds, 1982). Metacapnodium Spegazzini and
the t
Ophiocapnocoma Batista and Ciferri, wo genera of the
amily Metacapnodiaceae Hughes and Corlett apud Hughes
(Hughes, 1972), have been synonymized with Limacinia Neger
(Reynolds, 1985a).
Hypothesis: The Capnodiaceae sensu lato, including the
genera Acrogenotheca, Antennulariella, Capnodiun,
Euantennaria, Limacinia, Limacinula, Phragmocapnias,
Strigopodia, Trichomerium, Trichopeltheca, and Scorias,
comprise a monophyletic group.
B. METHODS.
1. Phylogenetic inference.
Compatiblity assisted parsimony is the strategy used in
this critique of the Capnodiaceae sensu lato. The aim of
this analysis is to achieve a parsimonious tree in the
manner suggested by (Wagner,1984; Farris and Kluge, 1986).
A cladogram is constructed on the basis of "good" char-
acter states, that is, character states that arise only
once. These characters are the compatible characters of
Meacham (1981); they are used to construct a convex char-
acter state tree. The noncompatible characters identified
are then accommodated to the basic skeleton to show
secondary cladistic relationships. Two steps are used.
Step 1. Character compatability analysis.
A history of compatibility methods in systematics was
reviewed by Meacham and Estabrook (1985). Opposing points
of view on this technique have been expressed -- often
acrimoniously -- by Duncan (1984, 1986), Churchill et al.
(1985), and Farris and Kluge (1985; 1986). No phylogenetic
reconstruction made with this method has been found in
mycological literature.
The basis of character compatibility analysis is that a
character or compatible groups (cliques) of character are
hypotheses of evolutionary relationships. Pairs of the 19
characters of the data set of this study were manually
tested for compatibility (Meacham, 1981) in order to dis-
cover cliques containing the largest number of characters.
The level of internal consistency was measured by the
ratio of the number of characters in the largest clique to
the number of characters in the data set; the fraction
approaching unity reflects fewest "false characters".
oo
Duncan (1980), Meacham (1980) and Meacham and Estabrook
(1985) emphasized the criterion of "convexity" for an
agreement between classification and estimated phylo-
genetic history. A character state tree asserts that its
states are convex. Each character state representing an
EU or EUs in a fully convex phylogenetic tree is uniquely
connected to a single immediately ancestral state by a
line representing some speciation event. According to
Meacham and Estabrook (1985), the convexity is demon-
strated, "if from any [EU] in the study, any of the other
species can be reached by passing along line segments of
the phylogenetic tree that pass only through other [EUs]
in that same collection. When a [character state tree]
asserts that its states are convex, it asserts that the
EUs] in a state share their common property by virtue of
inheriting it without change from other species in the
same state, and that the common property arose during the
speciation of the state's most recent common ancestor."
(2) Parsimony analysis.
Phylogenetic analysis using parsimony as a major criterion
developed over the last decade (Eldredge and Cracraft,
1980; Wiley, 1981; Duncan and Stuessy, 1984). The use of
cladistics in systematics has become common practice among
zoologists and vascular plant taxonomists. Specialists in
ferns (Funk, 1984) and mosses (Churchill, 1984) have used
this approach. Hoog (1979) made an attempt at fungal
analysis with calculation of patristic distances of a
preconceived tree.
The most complete phylogenetic work published done in
mycology has been by basidiomycete specialists (Hgiland,
1983; Vilgalys, 1986). Phylogenetic work with ascomycetes
has been abstracted by Currah (1984), Boise (1985) and
Reynolds (1985b). Wheeler and Blackwell (1984) provided a
concise discussion of cladistical methods and terms using
a basic three-species myxomycete model. Eriksson (1981)
discouraged the use of cladistics because he supposed a
lack of appropriate characters in fungi, although this
excellent review of loculoascomycete families amply
demonstrates the potential for phylogenetic systematics.
Preliminary results of cladistical analysis of the
Capnodiaceae was used in a discussion on pleomorphy
(Reynolds, 1986).
The parsimony analysis was carried out with Version 2.3 of
Phylogenetic Analysis Using Parsimony (PAUP), authored by
D. L. Swofford (1984). Several statistics were calculated
for each tree. The f-VALUE (Farris, 1972) (=total
homoplasy of Farris's Wagner 78 program; Klug and Farris,
1969 and Farris, 1970) is the sum of the deviation of
path-length from Manhattan distances across all distinct
pairs of taxa; low fractions indicate the least homoplasy.
The CONSISTENCY INDEX (Kluge and Farris, 1969) is the
minimum range of character-state changes in the data
divided by the actual length of the tree; fractions close
380
to unity indicate little homoplasy in a tree. The LENGTH
of the tree is the sum of character-state changes -- this
is also called patristic distance -- along the tree
branches. The BRANCH FACTOR (Rodman et al., 1984) is the
number of internal branches supported by unique
synapomorphies divided by the total number of internal
branches.
The determination of character polarity for the study
group is discussed in detail in the Results section.
Midpoint rooting was used for one analysis. Ingroup
analysis was used an another analytical method.
2. Assumptions.
The following assumptions were made for this phylogenetic
analysis of the Capnodiaceae.
(1) The genera utilized in this study are sister taxa.
The Capnodiaceae sensu lato embraces all of the genera
distributed into several families and more than one order
by Hughes (1976). Eriksson (1981) emphasized a
monophyletic view with his grouping of Hughes's families
into a single suborder. An alternative view is that of
Barr (1979, 1983) who distributed the taxa of the
Capnodiaceae sensu lato in two subclasses, three orders,
and four families.
(2) The phylogeny of the teleomorphs is representative of
the evolution of the family.
The capnodiaceous teleomorphs and conidiogenous fungi are
often produced from similar appearing hyphal morphs. The
darkly pigmented hyphal growths form a mycelial network on
living plant surfaces. Several kinds of these hyphal
morphs have been described (Mendoza, 1925; Hughes, 1976).
The similar morphology of the hyphae subtending
teleomorphic and anamorphic reproductive structures has
been used as an argument for phylogentic relationships of
teleomorphs and anamorphs (Hughes, 1976). Few of the
capnodiaceous holomorphs submised with this method have
been satisfactorily demonstrated to be comprised of the
proposed teleomorphic and anamorphic states (Reynolds,
1982). I have produced an anamorph from ascospore derived
cultures for Capnodium walteri (unpublished data). The
developmental cycle of Scorias spongiosa closely monitored
in nature allows a cautious admission of pleomorphy
(Reynolds, 1979). A convincing demonstration of
pleomorphy data is unavailable for other capnodiaceous
species. For this reason, this phylogenetic study is done
with a teleocentric focus; the pleomorphy suggested by
Hughes (1976) is not considered.
2
Reynolds: Foliicolous ascomycetes Ae
sensu lato. The genera utilized as operational taxonomic
units (OTU) and the major data sources are: Capnodium
(Reynolds, 1978), Limacinula (Reynolds, 1971, 1975a),
Antennulariella (Hughes, 1976), Trichomerium (Reynolds,
1982), Scorias and Phragmocapnias (Reynolds, 1979),
Acrogenotheca (Hughes, 1967), Euantennaria (Hughes, 1974),
Tricks eltheca (Hughes, 1965) , Limacinia (Reynolds,
1985a), and Strigopodia (Hughes, 1968).
4.'' Characters.
Only morphological data were available. The discussion of
hazards in the choice of characters and coding by Rodman
et al. (1984) is applicable. Characters were selected on
the following bases: The largest number of characters was
sought for which an argument of homology seemed
reasonable. The characters enabled character-state coding
which discriminated between genera.
Wheeler and Blackwell (1984) pointed out that conflicts in
the data set are warnings that one or more characters need
critical reexamination. The characters all have binary
coding (Table 1).
eoeeoeoeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeee
TAXON CHARACTER
i Keg Ds Ea (oe Ua dW Baa
character # 2 SAL S678 QeiOMl VOY Seas SUPT SL
ACROGENOTHECA VNOmOs OP OL OeOmOL dono ror ONO sO)
ANTENNULARIELLA JOP OR OFF O kel O16. 0 OnelanOn OM Onead VOsOnLO
CAPNODIUM ee Om Ome Ole ler OmOOmen Om. sO OmOrO
EUANTENNARIA OROZOROTOMOR DOF OO) 0 707 07:07 09:00 070
LIMACINIA OOP OF OO 0170), 0 .0:50)50750'10 20.0 00
LIMACINULA LO MOM a1 Onion OMOnr an Otel Os On law OneOuw OucO
PHRAGMOCAPNIAS LOO Ol One Ov Onli O. Or Le Oo
SCORIAS rl ee OmOrm Oran Onel TOMO On 1.) a OROL OO
STRIGOPODIA OF OOF ONOF OF OOO 0F0) 0710 (OX, 02:07 108 0
TRICHOMERIUM LO OlmorOr Owls OnO1> Op Olea OF ssOLiO
TRICHOPELTHECA OLORONO, ORO FORO OF ONOn Ol O02 0) 105 0870
TABLE 1. Character x taxon data matrix for Capnodiaceae
taxa. See text under MATERIALS:characters fora
discussion of the numbered characters.
eeeoeeeeeeeeeeeeeeeeveeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeeee
382
The characters are defined as follows:
CHARACTER 1. Dimorphic mycelium: present 0, absent 1.
This character reflects the occurence of the copious
mycelium as distinctive lower and upper portions. The
basal portion is in contact with the living plant surface.
The hyphae of the basal portion form a network in
Euantennaria, Limacinia, and Strigopodia; the hyphae of
Trichopeltheca are parallel to each other. Extensions of
the basal portion continue as a three-dimensional aerial
network in which the ascocarps develop. The aerial
network of Euantennaria and Limacinia (Fig. 1A) is more
prolific than that of Strigopodia. The aerial mycelium is
severely reduced to individual upraised hyphae in
oe and the ascocarps are produced from the
asa ayer of parallel hyphae.
CHARACTER 2. Stalk: absent 0, present 1. The ascus-
bearing portion of the ascocarp is subtended by an upright
column of tissue (Fig. 2C).
CHARACTER 3. Fruit body sessile: absent 0, present 1.
The fruit body develops on a mycelium formed of a
reticulate network of hyphae which rests on the surface of
the substratum.
CHARACTER 4. Limacinuloid: absent 0, present 1.
Limacinula species have thin walled, somewhat hyaline
hyphae which extend from the outer surface of the ascocarp
wall into the general substratum (Reynolds, 1971).
Eriksson (1981) found similarity in this habit to that of
taxa in the Coccodiniaceae and Naetrocymbaceae. Hughes
(1976) referred to these hyphae as occurring in a "fringe"
around the ascocarp. The ascocarp illustrated in Theissen
and Sydow (1917) is atypical.
CHARACTER 5. Ascocarp wall thickness: wide 0, thin 1.
The ascocarp wall of some species has a greater width than
in others (Figs. 2). The thicker walls were more than 3-4
cells in depth.
CHARACTER 6. Hyphal cell shape: cylindrical 0, spherical
1. The cells comprising the hyphal strands are either
cylindrical or somewhat spherical.
CHARACTER 7. Periphysoids: absent 0, present 1. Pericg
physoids are sterile elements in the ascocarp centrum in
addition to others such as pseudoparaphyses and paraphyses
(Luttrell, 1965) (Fig. 2). Eriksson (1981) questioned my
report of periphysoids in Capnodium (Reynolds, 1978). I
Figure 1. Diagramatic representations of characters of
the Capnodiaceae. A. Apical ends of the aerial portion of
the dimorphic mycelium of Limacinia fernandeziana. B.
Mycelium of Acrogenotheca elegans. C. Section through
outer portion of stroma of Scorias spongiosa.
565
2
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384
have examined the herbarium material that he utilized for
his study (kindly provided by him) as well as other coll-
ections. My opinion remains unchanged. Corlett (1970)
misinterpreted the periphysoids as pseudoparaphyses
(Reynolds, 1985a). This character-state datum was diff-
icult to determine for Antennulariella and Acrogenotheca.
The fruit bodies are very small and sterile elements are
difficult to demonstrate if present. The lack of space in
the small hymenial cavity may reflect a developmental
absence of this structure, when it should be present ina
phylogenetic sense. To account for both possiblities, the
character was initially scored for all possible states.
The PAUP algorithm accounts for this scoring by using the
most parsimonious character state had it not been missing,
when the tree length is computed. The taxon's location on
the tree is determined in congruence with those characters
without missing data. The position of the unknowns on the
resultant tree assigned a coding of "1" to this character
in Antennulariella and Acrogenotheca.
CHARACTER 8. Ascospore pigmented, muriform: absent 0,
present 1. OTU with only pigmented muriform ascospores.
CHARACTER 9. Ascospore hyaline, traversely septate: absent
0, present 1. OTU with only hyaline traversely septate
ascospores.
CHARACTER 10. Ascospore hyaline, muriform: absent 0,
present 1. OTU with only hyaline muriform ascospores.
CHARACTER 11. Ascospore hyaline, one septate: absent 0,
present 1. OTU with ascospores that are initially one
septate in the mature ascus; later becoming multiseptate
and pigmented usually after ejaculation.
CHARACTER 12. Mycelial persistence: persistent 0, deciduous
1. Mycelium persistent or deciduous. Refers do the
tendency of a mycelium to persist throughout the year or to
Slough off at a predictable time (Reynolds, 1975b).
CHARACTER 13: Hyphae parallel: absent 0, present 1.
Hyphae on the surface of the substrate are parallel to each
other rather than reticulately disposed.
CHARACTER 14. Stroma: absent 0, present 1. The mycelium
forms a stromatal structure (Reynolds, 1979) (Fig. 1C).
CHARACTER 15. Hyphal appendage: present 0, absent 1. The
ascocarp is appendaged with hyphal strands which project
from the surface of the ascocarp as though certain
meristematic cells continue in a hyphal-type of growth
(Figs. 2A).
Figure 2. Diagramatic representations of the characters of
the Capnodiaceae. A. Fruit body of Limacinia
fernandezana. B. Fruit body of Limacinula anomala. C.
Stalked fruit body of Capnodium walterii.
385
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CHARACTER 16. Antennulariella habit: absent 0, present 1.
The ascocarp is attached to the cells of the aerial hyphae
from which it originated... '\In addition, long, cylindricaL
flexuous hyphae originate from the outer ascocarp wall and
mingle with the rest of the mycelium. This character is
not homologous with Character 4 and Character 15.
CHARACTER 17. Phragmocapnias appendage: absent 0, present
1. The ascocarp is appendaged with short, acute,
pigmented, often unicellular appendages.
CHARACTER 18: Trichomerium appendage: absent 0, present
1. The ascocarp is appendaged with elongate, stiff,
usually multiseptate, pigmented setae.
CHARACTER 19. Acrogenotheca habit: absent 0, present 1.
The mycelium in contact with the living plant surface is
very reduced. The aerial hyphae from which the ascocarps
originate are upright and branch at wide angles (Fig. 1B).
C. RESULTS.
1. Character compatability analysis.
The primary analysis for character compatibility revealed
two overlapping cliques (Fig. 3) which accounted for all
characters. Seventeen characters (1, 3-8, and 10-19) are
common to both cliques. The common group plus character 9
and 17 form a primary clique (Figs. 3A and 4). The common
group plus character 2 form a smaller clique (Fig. 3B).
The patterns of character distributions representing both
of these cliques agree on the placement of the study group
EUs (Evolutionary Unit = Operational Taxonomic Unit of the
parsimony analysis). Antennulariella Acrogenotheca,
Euantennaria and Trichopeltheca have similar positions in
both character state trees. Both trees show a branch with
Limacinula, Capnodium, Phragmocapnias, Scorias, and
Trichomerium, but in different relationships. The final
estimate of evolutionary history for the Capnodiaceae
sensu lato was created from the primary clique with
characters 9 and 17 because of the greater compatibility,
and the full resolution of the tree.
The final undirected estimate of phylogeny for the genera
of the study EUs (Fig. 4) shows compatible relationships
between groups of genera. This character state tree
eoeeeoeeeeeeeeeeeee eeosveeereeeveeeeeeeeeeeeeeeeeeeeeeeeeeeneeeeeee
Figure 3. Character compatibility analysis of the
Capnodiaceae. ACReACrogenotnecai ANT=Antennulariella;
CAP=Capnodium; EUA=Euantennaria; LII=Limacinia;
LIU=Limacinula; PHR=Phragmocapnias; SC=Scorias;
STR=Strigopodia; TRM=Trichomerium; TRP=Trichopeltheca.
Seventeen characters are common to both cliques. A. The
common group plus Characters 9 and 17 form a clique. B.
The common group plus Character 2 form a clique.
388
asserts that its states are convex. A comparison with the
distribution of genera in the classifications which recog-
nize a Capnodiaceae sensu stricto counter the convexity of
the character state tree for the Capnodiaceae sensu lato.
The distribution of some characters on the estimate of
phylogenetic history suggests evolutionary trends. Spore
septation is one such trend. The septation pattern is
generalized in the OTUs on one end of the trees (Fig. 4B).
The ascospores of species in the ee group range
from transverse to muriform. On e other end of the
trees each OTU has one type of spore septation. Antennu-
lariella and Acrogenotheca produce ascospores which have
only one septum at maturity in the ascus (Character 11).
The spore dispersal strategy incorporates delayed stages
of hyphal initial germination and growth so that septation
and pigmentation develop in steps, usually outside the
ascus. Limacinula and Capnodium produce muriform
ascospores which are pigmented in the latter (Characters 8
and 10). Scorias, Trichomerium, and Phragmocapnias
produce ascospores with three cross septa eee SD jis
Mycelial persistence is a character of five genera on one
end of the tree and deciduous mycelium is produced by the
genera on the other end (Character 12). The thickness of
the ascocarp wall shows a correlated trend (Character 5).
Figure 4. Phylogenetic estimate of the Capnodiaceae and
distributions of characters and classifications using
character compatibility analysis. Abbreviations for the
families are given in Fig. 3A. The final undirected
estimate of phylogeny for the genera of the Capnodiaceae.
B. Distribution of spore septation types; general
septation; one septate (Character 11); muriform septation
(Characters 8, 10); traversely septate (Character 9). C.
Distribution of deciduous vs. persistent types
(Character 12). D. Distribution of several congruent
characters: Dimorphic hyphae (Character 1); Ascocarp wall
thickness (Character 5); Hyphal appendage on ascocarp
(Character 15). E-G. Distribution of taxa in proposed
Classifications. Solid and broken lines indicate taxon
broundaries; ?=taxon incertis cedis. OTUs outside taxon
boundaries were not considered in classification. E.
Distribution of taxa proposed in this study. Three
subfamilies are contained ina single family. F.
Distribution of Hughes taxa. Six families are distributed
in two classes and several orders. G. Distribution of
Barr's taxa. Four families are distributed in two
suborders and three orders. The Operational Taxonomic
Units (=EUs of the character state tree) in the unrooted
trees appear as two patristically equidistant clades in
relationship to the midpoint. OTU pairs Euantennaria and
Trichopeltheca, Limacinia and Strigopodia, an
Acrogenotheca and Antennulariella form one group and
Limacimula, Capnodium, Scorias, Phragmocapnias and
Trichomerium form the other group.
389
390
2. Parsimony analysis.
Parsimony analysis was carried out using midpoint rooting.
The tree is rooted at the midpoint of the longest path
connecting any pair of taxa with this method. The resul-
tant unrooted trees are represented in Figs. 5A and 6A-D.
The statistics for the trees were Length= 20.0, Consis-
tency Index= 0.95, f-Value= 14, and Branch Factor= 0.875.
The three midrooted tree generated by PAUP differ in the
topology of only one of the equidistant clades. The
relationships of Antennulariella, Euantennaria,
Limacinia, and Strigopodia are represented in Fig. 6A-C.
Euantennaria and peo eeeees are sister taxa in all
three configurations and are subtended by the node with
character state HTU-1 (Table 2). The three versions result
from their forming a sister pair to Strigopodia (Fig.
6A), to Limacinia (Fig. 6B), and to the sister pair
aise and Limacinia (Fig. 6C); the character state
of subtending nodes are HUT-2 in all three cases.
Character 2 is the only homoplastic character. A reversal
occurs in the branch clade terminated by Trichomerium.
3. Character polarity and tree rooting.
Proposals for estimating evolutionary character polarity
include outgroup analysis, ingroup analysis, the onto-
genetic method, and the paleontological method (Stuessy
and Crisi, 1984). Donoghue and Cantino (1984) state that
outgroup analysis has found most acceptance among pract-
itioners of parsimony analysis. They note the difficulty
of the use of outgroup analysis in angiosperm groups due
to a lack of well-corroborated hypotheses of cladistic
relationships among higher taxa. Cryptogams such as fungi
are no exception to this observation. Agreement is
lacking concerning the ancestor of the bitunicate asco-
mycetes as well as the sequence of descendants within the
bitunicate ascomycete. More specificially given the study
group used here as the Capnodiaceae sensu lato, what are
the monophyletic subgroups?
Eriksson (1981) proposed a phylogenetic sequence for the
evolution of the bitunicate ascomycetes; i.e. primitive
unitunicate ascus with thickened upper end which had a
pore, ascus with rostrum-type of endotunica and a ring,
semifissitunicate ascus with the extension of the
endotunica but without separation of the two tunica. The
fissitunicate ascus and the unitunicate asci were derived
from this point. Hawksworth (1982) suggested that several
modifications of the semifissitunicate ascus independently
led to modern groups, i.e. the bitunicate ascus would have
evolved with the modification of the endotunica; the
Xylariaceae and Helotiales were derived by a loss of the
ectotunica; the Lecanorales lichenized ascomycetes arose
with a loss of the endoascus ability to extend during
spore ejaculation and the gain of an apical ring.
591
The fossil record of fungi (Tiffney and Barghoorn, 1974)
shows evidence that the fissitunicate ascus coevolved with
the angiosperm leaf. Eriksson's (1981) "clades" 80 - 92
consist mostly of fungi that occur only on living leaves
(Hansford, 1946), including the taxa included in the
Capnodiaceae sensu lato.
Aside from the general question of the ancestor of
bitunicate ascomycetes, the descent of specific loculo-
ascomycetes is unresolved. Barr (1979) stated that the
Leptopeltidaceae was the ancestor of both unitunicate and
bitunicate ascomycetes. This reasoning implies that part
of the unitunicate ascomycetes evolved from an ancestor
common to the bitunicates. Eriksson (1981) lists the
unitunicate fungi before the bitunicate ascomycetes
implying that all unitunicate ascomycetes are ancestral to
bitunicate species. Eriksson believed the ascus in the
Erysyphaceae and the Meliolaceae to be bitunicate, while
Barr indicates an ancestral position for these fungi in
the unitunicate ascomycetes. Barr accepted the conclus-
ions of Holm and Holm (1977) that the Leptopeltidiaceae
ascus is unitunicate. Hughes (1976) considered the ascus
of the Capnodiaceae genus Trichomerium (Reynolds, 1982) to
be unitunicate because of the modification of the
endotunica. Eriksson (1981) considered this same
character merely a variation of the bitunicate ascus wall.
The origin of particular unitunicate ascus groups by
simplification of the fissitunicate ascus is suggested by
several studies. Minter and Cannon (1984) found both
unitunicate and functionally bitunicate species occurring
in the same family. "It just happens that the
Rhytismataceae is a natural family which is rather
variable in ascus opening mechanisms. That variability is
likely to provide valuable insights into the relation
between individual genera within the family, but it should
not be used to scatter the members of this family
throughout the ascomycetes." (I strongly disagree with the
inclusion of two diverse ascus types in the same family.)
Crisci and Stussey, 1980) and Stevens (1980) provide
similar guidelines for determining evolutionary
directionality. Criteria were grouped as independent or
first-level and dependent or second-level. The dependent
criteria of Stevens differ from the first-level criteria
of Crisci and Stussey in Stevens's rejection of ingroup
analysis. Under the circumstances, this seems to be the
best way to hypothesize polarity of capnodiaceous
characters.
Figure 5. Most parsimonious trees for the Capnodiaceae.
Family name abbreviations are listed in Fig. 2. The
position of synapomorphies are indicated by Character
number (Table 1) in the internal and terminal branches.
The HTUs (Table 2) are shown at the nodes. A. Midrooted
tree. B. Tree rooted with the sister pair Euantennaria
and Trichopeltheca as the designated ancestors.
STR
EUA
EUA
TRP
13
TRP
STR
13
Lil
Lil
ANT
ANT
ACR
12
ACR
11
19
LIU
LIU
CAP
CAP
SCR
SCR
PHR
TRM
TRM
3
One of the independent or first level criteria is the
character state distribution or "common is primitive". A
hypothetical ancestor can be based on the highest ratio of
"oO" to "1" coding for each of the OTUs (Table 1) and
optimized HTUs (Table 2). The ratios of VOU UL hin cone
present study group vary from 19/0 for Strigopodia to 10/9
Lor Gi els ect (Table 1). Thus, the most common "0"
state can be taken as primitive. Strigopodia would be the
OTU used for character polarization.
This method of rooting is similar to the Lundberg (1972)
method of outgroup selection. The PAUP program carried
out Lundberg rooting by first seeking the shortest
unrooted tree for the ingroup taxa. The resulting tree is
then rooted at the position in which a hypothesized
ancestor containing plesiomorphic states for all
characters would be attached to the tree (Swofford, 1984)
-- the “outgroup node" of Maddison et al. (1964) ates
corresponds to the Lundberg node (=Strigopodia) for this
study group (Table 2).
character # 1
RiIU=%
HIU=2
HTU<3
HTU-4
HTU=5
MLU=G
HTU-7
HTU-8
PRRPRPRPROO
PrRROOCOO N
COD0D0000 Ww
COD0D0D000 Ff
PRPRERPRPROO Ww
Soco°cc0o =H
PRPRPERPRPO AI
SCOC0D0000 ow
rPFrROOOC0CO wo
SCOCD0D000 OF
CODDORFOO FPR
PRPRPRFOOOO NPR
COD00D000 WP
CODD0D00O FPR
PRPRPRPROO UP
COCD0D00D00 APF
FOOCDOO0O NIP
CODD0D000 OF
SCO000000 oF
TABLE 2. Character x Hypothetical Taxonomic Unit data
matrix, for Capnodiaceae taxa. See text under MATERIALS
characters for a discussion of the numbered characters.
cooeeeeeeeeeeeeeeeeeeeeeeeeeeee eee eee eevee we eee ewe eee ewe eee eo ©
Ontogenetic states is another first level or independent
indicator of the direction of character state convexity.
The trend represented by ascospore ontogeny is represented
in Fig. 4B. Spore ontogeny progresses from a variety of
septation states in the species of a Single genus to a
single type of septation being characteristic of all
species ina taxon. The transseptate to muriform asco-
spores in the species of Euantennaria and Strigopodia
might be interpreted to represent arrested stages of spore
development. The maturing spore is first divided into two
cells, then more transsepta and longisepta form and the
cell might elongate. The derived genera produce spores
representing a single stage in this developmental
sequence; Antennulariella -- two celled at maturity in the
ascus, becoming more developed after dispersal; Scorias =-
three transsepta; and Limacinula muriform spores. A
major point is that these genera are also distinguished by
more than this one synapomorphy. This concept of spore
394
ontogeny was the basis of combining Ophiocapnocoma with
Limacinia (Reynolds, 1985).
A second-level or dependent criterion is group trends in
these taxa. The mycelium habit shows major changes which
I believe represent evolutionary progression in the leaf
surface microhabitat. The OTUs of the EUA-TRP-STR-LII
branch of the phylogenetic tree (Fig. 4A) produce the
fruit body upon or within the hyphae of a loosely woven
thickened mycelial mat which forms basal and aerial
portions. The mycelium becomes specialized in the ANT-ACR
branch (Fig. 4A). The mycelium is primarily a network of
surface hyphae in the LIU-CAP-SCR-PHR-TRM branch (Fig. 3).
Meacham and Estabrook (1985) point out that characters
that are related to each other functionally,
developmentally, or morphologically will have a dependent
compatibility. Evolutionary trends would act on suites of
these characters because of this dependency. One suite of
characters might be attributed to the distribution ina
tropical or temperate climate. Persistent mycelium and
pigmented ascospores seem characteristic of temperate
climate species. Deciduous mycelium and hyaline spores
seem characteristic of tropical climate species. A
transition from one type of environment to the other might
explain the change of spore strategy from a genus with
variable septation to a genus with a single spore
septation type and the change from persistent and
temperate to deciduous and tropical habit.
Evolutionary pressure might be attributed to the
positioning of the ascocarp for the most beneficial spore
dispersal from the fissitunicate ascus. Only Scorias
produces glabrous ascocarps; yet the ascocarps are
elevated by stalks which are subtended by a dense mycelial
mat. The genera produced in the aerial portion of a
looser mycelial mat, such as Limacinia and Euantennaria
are in a sense intercalary. The appendages are hyphal
outgrowths. Antennulariella is usually found among the
aerlal mycelium of other capnodiaceous species; the
appendages are specialized hyphal outgrowths, differing in
morphology from the generative hyphae (Character 16). The
ascocarps of Limacinula produce a somewhat similar growth,
but in a different habitat (Character 4). Stalks subtend
the ascocarps of eee and Phragmocapnias (Character
2) to elevate them from e substratum.
Figure 6. Analysis of Cladogram. Abbreviations for the
family names are given in Fig. 3. A-C. Three
configurations of the midrooted tree (Fig. 5). OTUs and
HTUs are indicated as explained in Fig. 5. D. The_ |
plesions Euantennaria and Strigopodia are compressed into
the node with an identical character state. E-F. Tree
represented by clade in D with designated root.
OGN=Outgroup Node; IGN=Ingroup Node. E. Tree rooted
with plesion Strigopodia at IGN. F. Tree rooted with
plesion Euantennaria at IGN.
STR
EUA
TRP
Lil
Lil
STR
EUA
Lil
STR
Lil
EUA
595
HA
TRP
Lil
HA
Lil
396
Two roots are suggested using these methods of determining
character polarity. The OTU and HTU character state with
the commonest distribution, all "0", is that of Euanten-
naria and of HTU-1. Thus a hypothetical ancestor would be
coded with "0" for all nineteen characters in the data
matrix of this study group. The use of Euantennaria and
its sister taxon, Trichopeltheca, for character polar-
ization is in agreement with the position assigned these
genera to a terminal end of the unrooted tree; the tree
produced by the character compability analysis shows
Similar positions. The tree rooted in this manner shows
Character 7 as a plesiomorphy on the main stem. An alter-
native root would be the sister taxa Strigopodia and
Limacinia. These two OTUs are positioned in the same end
of trees generated by both analyses. The tree rooted in
this manner shows Character 7 as an apomorphy on a clade
terminated by Euantennaria and Trichopeltheca, rather than
the main stem.
Further support for a choice of an ingroup taxon for
characterization polarization is derived from consider-
ation of the plesion nature of Euantennaria and
Strigopodia. The character state of Euantennaria is the
same as HTU-1 (Tables 1 and 2). The character state of
Strigopodia and HTU-2 are the same. Thus, if the OTU is
suppressed" to the node with the same character state, it
assumes the position of an ancestor. The unrooted trees
bearing Strigopodia and Euantennaria then become identical
(Fig. 6D). Euantennaria is seen as the ancestoral taxon to
Trichopeltheca; Strigopodia is ancestral to Euantennaria
and to Limacinia.
Rooting with the plesion Strigopodia (Fig. 6E) as the
Ingroup Node (Maddison et al., 1984) produces an
unresolved tree. A tricotomy results from Euantennaria
and Limacinia. Furthermore, Trichopeltheca is shown as a
monocotomy. Rooting with the plesion Euantennaria as the
Ingroup Node (Fig. 6F) produces a resolved tree.
Trichopeltheca is descended from Euantennaria at the
Ingroup Node distinguished by autapomorphy Character 13.
Sa aoe is descended from Euantennaria with the change
in Character 7; Limacinia is descended from Strigopodia
and is distinguished by the autapomorphy Character 6.
D. CONCLUSIONS.
The trees produced in this study support the hypothesis
that the Capnodiaceae sensu lato is monophyletic. Several
subclades, recognized proposed here as subfamilies,
generally correspond to taxa proposed by others.
Congruence with the Barr and Hughes classifications 1s
achieved with the recognition of the phylogenetic |
relatedness of the study group, rather than scattering the
genera through several suprafamilial taxa. The scheme for
Ascomycetes proposed by Eriksson (1982) is the acceptable
ordinal framework for the classification.
og?
I recognize the following taxonomic treatment of the
Capnodiaceae.
Subclass Euascomycetidae
Group Ascoloculares
Order Dothideales
Suborder Capnodioideae
FAMILY Capnodiaceae (Saccardo) Hohnel, pro parte,
teleomorphoses only.
Hohnel (1910); Spegazzini (1918), Theissen and Sydow
(1917), Arnaud (1911), Batista and Ciferri (1963),
Hughes (1976), Reynolds, (1978); Eriksson (1981).
Basionym=subfamily Capnodiaceae Saccardo
Saccardo (1880)
=Antennulariellaceae Woronochin, pro parte,
teleomorphoses only. Woronochin (1925); Hughes
(1976).
=Euantennariaceae Hughes & Corlett in Hughes, pro
parte, teleomorphoses only. Hughes (1972); Hughes
(1976).
=Metacapnodiaceae Hughes & Corlett in Hughes, pro
parte, teleomorphoses only. Hughes (1972); Hughes
(1976).
=Triposporiopsidaceae Hughes, pro parte,
teleomorphoses only. Hughes (1976).
SUBFAMILY Capnodioideae Saccardo
Saccardo (1882)
Type genus= Capnodium Montagne: Montagne (1849);
Reynolds, (1978).
This provisionally recognized established subfamily
includes the following genera:
Limacinula (Saccardo) Hohnel: Hohnel (1909);
Reynolds (1971).
Phragmocapnias Theissen & Sydow: Theissen and Sydow
(1917); Reynolds (1978).
Scorias Fries: Fries (1825); Reynolds (1979).
Trichomerium Spegazzini: Spegazzini (1918);
Reynolds (1982).
SUBFAMILY Limacinioideae subfamily novo prov.
Type genus= Limacinia Neger in Johow: Johow
(1896); Reynolds (1985).
398
This provisional subfamily also includes the following genera:
Euantennaria Spegazzini, pro parte, teleomorphosis
only; Spezazzini (1918); Hughes (1975, 1976).
Strigopodia Batista in Batista, Maia, & Vital, pro
parte, teleomorphosis only: Batista, Maia and
Vital (1957); Hughes (1968).
Trichopeltheca Batista, Costa & Ciferri, pro parte,
teleomorphosis only: Batista, Costa and
Ciferri (1957); Hughes (1965), Corlett (1970),
Corlett etal. (1973).
SUBFAMILY Antennularielloideae subfamily novo prov.
Type genus= Antennulariella Woronochin, pro parte,
telemorphoses only: Woronochin (1915); Hughes
(1976).
This provisional subfamily also includes the following genus:
Acrogenotheca Ciferri & Batista in Batista & Ciferri,
pro parte, telemorphoses only: Batista and
Ciferri (1963); Hughes (1967, 1976)
=Brooksia Hansford, pro parte, teleomorphosis only:
Hansford, 1956; Deighton & Pirozynski, 1966).
Cooke and Hawksworth (1970) assumed that Hohnel (1910)
referred to Saccardo (1882) when they recognized the
Capnodiaceae (Sacc.) Hohnel with the basionym as the
Saccardo subfamily Capnodiaceae in his Fragmente zur
Mykologie 532 entitled, "Ubersich der Capnodiaceen-
gatungen"; later reference was made to the Capnodiacee
(Hohnel, 1911). The authorship by F. von Hohnel was
recognized by Hughes (1976) as per Cooke and Hawksworth
(1970), and by Barr (1979). Arx and Muller only used
Hohnel with no reference to Saccardo. Eriksson (1981,
1982) attributed the family name to Theissen and Sydow
(1917); these authors recognize Hohnel (1910) as the
author of Capnodiaceae and give reference to Saccardo
(1910).
The ideal way to compare the results of this study with
other phylogenetic hypotheses to compare representative
trees. The classifications of Hughes and Barr provide
hypotheses for comparative purposes. Devising a tree to
represent these views requires the construction of tree
topologies to representing their implied phylogenetic
hypothesis. Another approach is to impose the
classification of these authors on the trees produced in
this study.
The character state tree for Hughes's (1976) four family
classification in represented in Fig. 4F. The polyphyly is
created by suprafamilial gerrymandering. Limacinia was
599
assigned to the order Pleosporales (Corlett et al. 1973).
Limacinula was suggested as a member of the order Chae-
tothyriales. Trichomerium was said to have a unitunicate
ascus and therefore in a different subclass. The convex-
ity is restored by ignoring these peccadillos by bringing
Limacinia, Limacinula, and Trichomerium into more direct
lines of descent with other capnodiaceous taxa.
The character state tree for Barr's (1979) classification
of the study EUs as they would appear in the orders of the
Loculoascomycetes is represented in Fig. 4G.
Antennulariella, Acrogenotheca and Trichomerium were not
mentioned. The remaining capnodiaceous genera are
seperated from each other by grouping with other taxa in
three orders and four families. Gross polyphyly is thus
introduced into the evolutionary history of capnodiaceous
genera with the filial distance imposed by Barr's
phylogenetic hypothesis.
The clades of the rooted tree from the parsimony analysis
can be compared with the classification proposed by Hughes
(1976). His Euantennariaceae corresponds to the most
ancestral clade with the sister taxa Euantennaria and
Trichopeltheca. The Metacapnodiaceae, containing the
genus Limacinia (fide Reynolds, 1985a) is a clade;
Strigopodia, a taxon of uncertain affinity in the Hughes
system, is a sister taxon. Antennulariella, the taxon in
Hughes's Antennulariellaceae, and a sister taxon,
Acrogenotheca are the terminal taxa of a clade.
Limacinula was of uncertain affinity; it the terminal
taxon of a branch. The three sequential taxa, Capnodium,
Scorias, and Phragmocapnias were placed in the
Capnodiaceae sensu stricto by Hughes. The last taxon on
the tree depicted in Fig. 4F, Trichomerium, was said by
Hughes to produce a unitunicate ascus and was banished
from the loculoascomycetes on this basis.
The data from the parsimony analysis are uncorrelated with
the Barr (1979, 1983) classification. The nine OTUs of
this study to be found in her keys were distributed by her
in two subclasses, three orders, and four families.
Euantennaria and Trichopeltheca would be assigned to the
Subclass Loculoparenchymatomycetidae, Order Dothideales,
Family Euantennariaceae. The subclass
Loculoedaphomycetidae would contain Limacinia (fide
Reynolds 1985a) of the Family Metacapnodiaceae and
Strigopodia and Limacinula of the Family Naetrocymbaceae
of ene Order Chaetothyriales. Capnodium, Scorias, and
Phragmocapnias were placed in the Order Asterinales,
Capnodiaceae (sensu stricto).
The Capnodiaceae recognized here can serve as the outgroup
in the universally parsimonious construction of the
phylogeny of other loculoascomycete groups. Maddison et
al. (1984) note in their discussion of outgroup analysis
and parsimony that parsimony in the outgroups is highly
desirable. Accordingly, the capnodiaceous might be used
as an outgroup for analyses that examining relationships
400
within the Chaetothyriaceae Hansford ex Barr (1979) and
the Herpotrichiellaceae Munk (1957).
VI. BIBLIOGRAPHY.
ARNAUD, G. 1911. Contribution a 1! étude des fumagines.
2% Systematique et organization des espéces. Ann. Ecole
Natl. Agric. Montpellier, sér. 2, 10:211-330. ARX, J. A.
von, and E. MULLER. 1975. A re-evaluation of the
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MYCOTAXON
Vols XXVET, pp. 404 “October-December 1986
ihe ih Misti isos LUCE adn nto Aco a e bole taa tat i eh wicks bal f°
NiO Tit eee
Announcing a
GUIDE TO THE TREATMENT OF SANCTIONED FUNGAL NAMES
Subcommittee C of the IAPT Committee for Fungi and Lichens has
elaborated a Report and a set of Proposals to alter the I1.C.B.N.,
in order to clarify the situation of sanctioned fungal names
(publication scheduled in Taxon, Nov. 1986).
A preliminary version of a Guide to the treatment of sanc-
tioned names is ready, but still contains a few alternative solu-
tions, on which decisions by the IAPT Committee for Fungi and
Lichens and the next Botanical Congress are pending. After the
Committee has given its vote, a modified, but still preliminary,
version of this guide is to be published in Mycotaxon. Until then
photocopies of the text are available upon request from Dr. W. Gams,
Centraalbureau voor Schimmelcultures, P.0.Box 273, 3740 AG BAARN,
Netherlands.
MYCOTAXON
ea et en et
Vol. XXVII, pp. 405-449 October-December 1986
THE USE OF ISOZYME ANALYSIS IN FUNGAL
TAXONOMY AND GENETICS1; 2
J. A. MICALES3
Department of Plant Pathology
Cornell University
Ithaca, NY 14853
M. R. BONDE and G. L. PETERSON
Foreign Disease-Weed Science Research Unit
Building 1301, Fort Detrick
Frederick, MD 21701
SUMMARY
| Isozyme analysis, as performed with starch
: gel electrophoresis, is a valuable tool in the
study of fungal taxonomy and genetics. Appli-
cations and procedures for this technique are
presented and methods of data interpretation
are discussed. It is recommended that in most
studies a systematic approach be used whereby
the presence of many specific enzymes is tested
and the best buffer for their resolution is
determined in an initial screen. This even-
IThis study was carried out at Frederick, MD and partly
funded by the United States-Israel Binational Agricul-
tural Research and Development Fund Grant No. US-563-82.
2Mention of a trademark or propriety product does not
constitute a guarantee or warranty of the product by USDA
and does not imply its approval to the exclusion of other
products that may also be suitable.
3current address: USDA, U. S. Forest Service
Forest Products Laboratory
Box 3150
Madison, WI 53705
406
tually allows the greatest amount of
useful information to be obtained with
the least amount of effort when larger
numbers of isolates are later examined.
INTRODUCTION
Isozyme analysis, as performed with starch gel
electrophoresis, is a versatile and inexpensive technique
which can be used to determine the amount of genetic
variation among organisms. This procedure is commonly
used in population genetic studies of fish (1, 19, 23),
mammals (33, 42), insects (43), nematodes (16), and
plants (9), but its application to fungal taxonomy and
genetics is just being realized. The objective of this
paper is to acquaint the reader with the theory of
isozyme analysis and to provide detailed procedures so
that the technique can be more widely applied. Reviews
of this topic have been published (10, 41), but these
have been written for plant and animal geneticists. This
paper will stress the application of isozyme analysis for
mycologists and plant pathologists and will address
problems inherent to working with fungi.
Enzymes which are coded by different alleles or
separate genetic loci frequently possess different
electrophoretic mobilities. Such differences are due to
variations in the amino acid content of the molecule,
which in turn is dependent on the sequence of nucleotides
in the DNA. Electrophoretic banding patterns are
frequently predictable since they are dependent on the
genetic and nuclear condition of the organism. Most
mycologists and plant pathologists who use _ electro-
phoresis do not take advantage of this relationship and
restrict their interpretation of electrophoretic data to
simple "band-counting” techniques (34, 36); a genetic
interpretation of the same data frequently provides more
information about the genetics and taxonomy of a group of
organisms. Genetic crosses may be necessary to confirm
genetic interpretations, but certain banding patterns are
readily recognizable from comparable studies in human and
animal genetics (15). Much interest has been expressed
in this technique, and it is currently being used in our
laboratory to study the taxonomy and genetics of fungal
taxa.
407
APPLICATIONS OF ISOZYME ANALYSIS
The ambiguous results of early studies of isozyme
analysis with fungi (36) originally discouraged the
application of this technique in mycology and plant
pathology. More recently, isozyme analysis has been used
to quantify genetic variation in Agaricus (21, 22, 31,
32), Entomophthora (20), Neurospora (38), Peronosclero-
spora (3), Phytophthora (40), and Puccinia (5, 6). The
potential applications of isozyme analysis with fungi are
limited only by the imagination of the researcher. There
are at least three major areas in which isozyme analysis
can be used. These include the clarification and
delineation of fungal taxa, the identification of fungal
cultures to the species or subspecies level, and the
study of the genetics, including population genetics, of
specific fungi. A list of representative fungi used for
isozyme analysis, and enzymes found to give interpretable
results, is presented in Appendix I.
Clarification and delineation of taxa -_ The
interpretation of banding patterns in terms of specific
alleles allows the determination of ratios of alleles
expressed in common among fungal isolates; these ratios
are an excellent means of determining phylogenetic
relationships among organisms. In studies of closely
related species, the assumption is made that the same
loci are present and are being compared among isolates,
even though the organisms may be from different species
and the number of chromosomes may not be the same.
Isozyme analysis can be very helpful in solving
taxonomic problems when there are few morphological
parameters or when other characteristics are very plastic
among isolates of the same species. It is relatively
easy to detect 15 or more enzymes that can be used to
differentiate and delineate closely related species.
Examples of the use of isozyme analysis for taxonomic
clarification have been presented by Royse and May (31,
32), Bonde et al. (3) and Micales et al. (24, 25).
Identification - Up to 90 different enzymes can be
easily tested to determine whether a particular species
can be distinguished from other species by means of the
presently available techniques of isozyme analysis. Many
of these enzymes will not be suited for a given fungal
408
species. In examining the smut pathogen Tilletia indica
Mit., Bonde et al. (2) readily found 29 enzymes of the 48
tested that were in high enough concentrations to warrant
further studies.
With any species, some genetic loci tend to have
little or no variation among individuals. These loci are
termed “monomorphic” and are defined as loci in which 997%
or more of the individuals have the most common allele.
Although it is premature to estimate the average
percentage of monomorphic loci for fungi in general, the
work of Bonde et al. (2) suggests that about 50% of the
genetic loci of T. indica are monomorphic. Tooley et al.
(40) found that 11 of the 24 loci tested were monomorphic
in Phytophthora infestans (Mont.) deBary. It is these
monomorphic loci which perhaps should be stressed when
comparing an "unknown" isolate to previously identified
cultures in order to determine identity. Studies with
other fungi in our laboratory suggest that some fungal
species have considerably less allelic variation.
The importance of the correct identification of
pathogens cannot be overstated. For example, regulatory
agencies at the federal and state levels in the United
States must correctly identify pathogens of regulatory
significance. In some cases, all a regulatory official
has to work with are a few spores in a shipment of
produce; no symptoms are available. These spores may
have been picked up from weeds during harvest; the
identity of pathogens on weeds is often unknown. Isozyme
analysis is a tool which can facilitate identification.
Fungal genetics - Isozyme analysis also is being
used in fungal genetics. Genetic studies rely on loci
which exhibit allelic variation. Tooley et al. (40) were
able to demonstrate the probable random mating patterns
of Phytophthora infestans in Mexico where the sexual
stage of the pathogen exists; this was done by examining
the genotypic distributions at two polymorphic loci.
Almost every individual in the asexual population,
representing the United States, Canada, and Europe, was
heterozygous at these two loci, thus demonstrating the
mear absence of mating. Random mating also was
demonstrated among sexual populations oof Puccinia
graminis f. sp. tritici by Burdon and Roelfs (5, 6).
Most studies have not been able to correlate specific
409
isozyme patterns with differences in pathogenicity. One
exception was reported by Burdon and Roelfs (5), who were
able to find such a correlation in asexual populations of
Puccinia graminis f. sp. tritici. This association was
not true in sexual populations which had a much higher
degree of genetic variability. Isozyme patterns of
Puccinia recondita have been used to trace independent
introductions of the pathogen to the United States (6).
The potential for isozyme analysis in such studies of
fungal genetics is great; the technique will be more
widely used in the future as additional genetic markers
are reported.
SELECTION OF FUNGAL TISSUES
Electrophoresis has been performed successfully with
a wide variety of fungal tissues, including vegetative
mycelia, conidia, basidiocarps, and sclerotia. Care must
be taken in the selection of fungal tissues’ since
inconsistent results may be obtained when a mixture of
different tissues is used. Such variation may be due to
the differential expression of alleles at different
stages of development or from variations in the nuclear
condition, as in the expression of the monokaryotic and
dikaryotic stages of the organism. Variations in
catalase banding patterns of conidia and vegetative
mycelia have been reported for Fusarium solani (Mart.)
Appel and Wr. (14). Similar differences have been
observed among different portions of the basidiocarp in
Lentinus edodes (Berk.) Sing. (2/), Coprinus cinerens
(Schaeff.) Cke. (26), and Agaricus bisporus (Lange)
Imbach (28). The importance of using uniform tissue
samples is demonstrated by such experiments.
SAMPLE PREPARATION
Proteins can be extracted from spores, mycelia, or
fruiting bodies which have been grown in culture or
collected in the field. Uniform conditions of light,
temperature, and nutrition should be used when sample
material is grown in culture in order to minimize the
chance for the differential expression of induced
enzymes. Ideally, samples should be collected at a
specific stage of the growth curve; this is usually
difficult to coordinate, and most researchers harvest
their material after a specified period of growth.
Initial studies should be conducted to determine the
410
amount of electrophoretic variation detected at different
stages of the growth cycle. The amount of protein in the
mycelium may also vary with the stage of growth; protein
assays (4, 17) should be conducted to determine the time
of maximum protein production.
Proteins can be extracted from fungal tissues by a
number of different procedures. Thin-walled structures,
such as certain conidia and vegetative tissues, can be
crushed with a glass rod after the sample has been frozen
in liquid nitrogen. lLyophilization is frequently used
with vegetative tissues and large fruiting bodies; this
process removes the water from the sample and allows it
to be readily crushed. The preparation of acetone
powders can be used in lieu of lyophilization although we
have obtained lesser quantities of enzymes with this
technique. Large fungal structures, such as_ basidio-
carps, ascocarps, or mycelial mats, can be disrupted with
a variety of commercial homogenizers or simply crushed in
a chilled mortar and pestle. Sample buffer is added to a
freeze-dried sample before the sample begins to thaw or
rehydrate. Non-frozen samples must be disrupted in a
buffer solution. The amount of buffer to be added will
vary with the nature and the size of the sample and is
best determined empirically; there is a thin line between
the addition of enough buffer for adequate sample size
and diluting the sample below the level of detection.
Proteins in the sample may precipitate if insufficient
buffer is added or if the buffer is of the incorrect
ionic strength.
The denaturation of enzymes must be prevented during
and after homogenization; samples should be kept below 4°
C during preparation and storage. Agents which inacti-
vate proteases, such as polyvinylpyrrolidone (PVP), may
be added to the sample buffer to prevent proteolysis.
PREPARATION OF STARCH GELS
The preparation of starch gels is something of an art
and requires some practice in order to obtain repro-
ducible electrophoretic patterns. Many different
electrophoretic systems are available commercially, but
many researchers construct their own equipment with the
inexpensive materials described below. Modifications of
this system are possible if one needs to vary the size
and depth of the desired gel. Diagrams of electro-
411
phoretic systems are presented by Conkle et al. (10),
Werth (41), and May (19) and illustrated in photographs
in Fig. 1.
Two types of electrophoretic experiments are usually
distinguished. A "screening" run should be conducted
early in a study with a few isolates (5-10) and a large
number of stains and buffer systems. This experiment
will determine which enzymes are active in the taxon and
which buffers provide optimal resolution of each enzyme.
The taxonomy and genetics of large numbers of isolates
can then be studied in subsequent “analytical” runs,
using the enzyme systems determined to be best in the
screening run. The nature of the experiment (screen vs.
analytical) will influence the preparation, sample
application, and final handling of the gel; these
differences will be detailed in the following sections.
Gel molds - Gel molds can be prepared by clamping
plexiglass strips (163 x 20 x 6 mm and 215 x 20 x 6 mm)
by means of binder clips around the four edges of a piece
of plate glass (254 x 164 x 5 mm) as shown in Fig. l.
The thickness of the gel can be increased for compre-
hensive screening runs to test for large numbers of
enzymes. This can be done by using thicker plexiglass
strips or by securely taping two strips together. The
mold should be placed on a level surface; uneven gel
thickness will cause variations in enzyme migration rates
in different regions of the gel.
Gel preparation - Gels can be prepared with 42 g of
refined potato starch (Sigma Co., St. Louis, MO) per 300
ml gel buffer. Formulas for commonly used buffer systems
are presented in Appendix II. Thicker gels for screening
runs can be prepared with 70 g of starch in 500 ml of gel
buffer. The starch is weighed and placed in a 1000 ml
wide-mouthed flask (Fig. 1); approximately 1/3 of the gel
buffer is added and the suspension vigorously swirled
until the suspension is free of lumps. The flask can be
placed on a stir plate at this time; the suspension
should be continuously stirred to prevent settling of the
starch. The remaining gel buffer is heated to boiling
and rapidly poured into the flask which contains the
starch. Care must be taken to prevent burns on the hands
and face from the escaping steam; the boiling gel buffer
should be initially swirled to allow the steam to
escape. The hot suspension must be vigorously swirled
412
413
until it is uniformly mixed; the starch should _ be
discarded if any lumps are visible since they will not be
removed by continued mixing. The flask is then placed
over a Bunsen burner and heated until boiling. The
suspension must be swirled occasionally to prevent the
starch from burning on the bottom of the flask. The
suspension should be heated until it is boiling rapidly;
it will also lose some of its viscosity and become
somewhat transparent as the starch goes into solution.
The flask then is removed from the flame, and the starch
solution is degassed for approximately 20 sec to remove
air bubbles. This usually is done with a vacuum formed
by a water aspirator; a vacuum pump can also be used. A
rubber stopper with two glass tubes is placed in the
mouth of the flask. The hose attached to the vacuum
source is placed over one glass tube; the other is
covered with a finger tip so that the operator has direct
control over the operation of the vacuum. A trap should
also be inserted into the line to prevent the backflow of
water from the aspirator into the gel. The length of
time of degassing will vary with the volume of starch and
the type of gel buffer; the vacuum should be applied
until all small bubbles are removed from the solution and
large bubbles are distributed uniformly throughout the
mixture. The vacuum should be removed smoothly from the
flask to prevent the formation of additional air
bubbles. The solution is then poured carefully (Fig. 1)
into the gel mold, and bubbles and small foreign
particles can be removed before the starch cools and
solidifies. The gels should be allowed to cool for at
least one hour before electrophoresis. Gels should be
covered with plastic wrap upon cooling to prevent loss of
moisture. The plastic wrap should be spread as smoothly
as possible, as wrinkles will cause depressions in the
gel surface.
Fig. 1. Preparation of starch gels for electrophoresis.
A) Starch is added to and dissolved in hot gel-buffer.
B) Air is removed under vacuum and C) poured into a gel
mold. D) Paper wicks with sample extracts are placed at
the origin. E) Starch gel with 4 repeat series of sample
extracts. After electrophoresis, gel will be cut
vertically through lanes with marker dye and _ sliced
horizontally to produce 8-10 thin layers. F) Starch-gel
apparatus during electrophoresis run. Blue Icd®) keeps
gel cool.
414
APPLICATION OF SAMPLES TO THE GEL
The plastic wrap is removed from the gel surface
after the samples have been prepared. A scalpel is used
to cut around the periphery of the gel; the binder clips
and long plexiglass strips can be removed at this time.
Excess starch is trimmed from the surface of the shorter
plexiglass strips. The origin of the gel is formed by
making a cut across the length of the gel approximately
2.5-3.0 cm from the cathodal edge (Fig. 1). The cut must
be perpendicular to the gel surface and reach the bottom
of the gel. The cathodal strip is pulled away from the
cut surface; this facilitates the application of samples
to the anodal edge of the origin.
Small filter paper wicks are used to apply the
protein to the gel (Fig. 1). Wicks can be cut from large
filter paper sheets or strips; a variety of filter paper
grades can be used, but thick grades will waste precious
sample material. Whatman No. 3 filter paper is used in
our laboratory. The size of the rectangular wicks varies
with the number of samples that are to be placed on the
gel. We usually use wicks that are 2-3 x 12-15 mm;
longer wicks are required for thicker screening gels.
Wicks are dipped into the crude homogenate and
removed after the sample has risen by capillary action to
within 1/3 of the top. Do not allow the wick to touch
solid fungal material since this will cause streaking in
the gel. The wick is then placed vertically on the
anodal surface of the origin; the bottom of the wick
should be flush with the bottom of the gel. Samples
should be spaced with approximately 1 mm between each
wick (Fig. 1); up to forty wicks can be applied to a gel
in this manner. Larger spaces can be used at certain
intervals to facilitate the determination of the location
of each sample on the gel after staining. Food coloring
is diluted with sample buffer (1:1) and applied to two
wicks; these dye markers are placed at each end of the
gel so the extent of migration during and after
electrophoresis can be monitored.
The arrangement of samples on the gel depends on the
type of experiment. In a screening run, we generally use
a sequence of 8-10 samples repeated four times (Fig. 1).
Dye markers are placed between each group at the origin
to facilitate determination of where to cut the gel
415
following electrophoresis. A larger number of samples is
used for analytical experiments. The sequence of samples
should be repeated at least once in an analytical run if
sufficient space is available; variations in migration in
different portions of the gel can then be detected and
this greatly facilitates interpretation of results. It
is advisable to repeat one or two samples at intervals on
the gel; this provides a standard to which each sample
can be compared. If particular comparisons are desired
among isolates, these samples should be placed in close
proximity to insure accurate analysis.
ELECTROPHORESIS
The cathodal strip is moved back into place after the
wicks have been applied (Fig. 1). The two pieces of gel
must make firm contact in order to allow the electrode
buffer to move freely between them. The surface of the
gel is then covered with plastic wrap which is folded
back 1.5 cm from the anodal and cathodal edges of the
gel. Buffer trays (we styled them from the tops of
plastic butter dishes) are filled with the appropriate
electrode buffer; absorbant, disposible cloths (e.g.
Handiwipes®) are placed in these trays. These cloths
should be folded into three layers and thoroughly washed
of all detergent before use. The leading edge of the
cloth is placed on the surface of the gel making certain
that there is good contact between the cloth and the
gel. The remaining plastic wrap is folded back over the
surface of the absorbant cloth. Platinum electrode
wires, which are soldered onto the end of alligator clips
attached to a DC power supply, are inserted into each
electrode buffer tray, with the cathode (negatively
charged, black lead) closest to the gel origin (Fig. 1).
Power is supplied to the gel and adjustments are made so
that the levels of voltage and current are appropriate.
Proteins are allowed to migrate out of the wicks; this
process takes about 15 min and can be estimated by the
rate of migration of the tracker dye. After this time
interval, the power is turned off, the absorbant cloth on
the cathodal end is lifted, and the wicks are removed.
The two slices of gel are firmly pushed together, and the
absorbant cloth and plastic wrap reapplied. The gels
must be kept cool during the electrophoretic run. We use
Blue Ic packs on top of two layers of 5 mm-thick
plate glass placed directly on the surface of the gel to
insure uniform cooling (Fig. 1); commercial cooling
416
devices also are available. Power is resupplied; the
levels of current and voltage are readjusted and
monitored throughout the electrophoretic run. A typical
experiment requires approximately 2.5-3.0 hr of migration
for adequate resolution of protein bands, although some
researchers use lower voltage and current settings and
allow the gel to run for longer periods of time.
A list of gel buffer systems commonly used in our
laboratory is presented in Appendix II.
SLICING THE GELS
After electrophoresis is completed, the power supply
is turned off, and the plastic wrap, absorbant cloths,
buffer trays, and plexiglass strips are removed from the
gel. The anodal strip above the dye markers is cut away,
and an identifying mark is cut into the anodal and
cathodal gels in a _ standardized position for proper
orientation of the gel after staining. One advantage of
starch gel electrophoresis is that several’ thin,
horizontal slices can be made from each gel; these can be
stained individually for the detection of different
enzymes. Commercial gel slicers are available, but many
researchers use plexiglass guides (300 x 25 x 1 mm) and
nylon sewing thread (“invisible thread") or low test
fishing line. Slices are cut by sequentially stacking
the plexiglass guides along the long edges of the gel and
by drawing the thread through the gel over the surface of
the guides (Fig. 2). The uppermost slice is usually
discarded due to the thick "skin" which is formed during
cooling; electrophoretic migration is frequently
inconsistent in this uppermost region of the gel. The
remaining slices are gently removed from the gel and
placed in styrofoam staining trays; the side of the slice
which was closest to the center of the gel should be
placed upright since enzyme concentration is highest in
this region.
Enzyme screens require many more duplicate gel pieces
than do analytical runs, thus allowing the testing for
the presence and resolution of a large number of enzymes
from a single gel. The procedure used for cutting gels
for enzyme screens is similar to that described above,
except that vertical cuts must also be made through dye
markers to separate the individual groups of samples.
This is usually done after the horizontal slicing has
ae LOI SO
417
been accomplished. A distinguishing cut or notch must be
placed in each section of the gel so that the orientation
of samples and identification of the gel buffer system
can be maintained during later staining and manipulation
of the many gel pieces. One section of each gel is then
placed in each staining tray or on individual glass
plates, the latter for agar overlays, fluorochromes, and
certain azo dyes (see Appendix III), so that the
different gel buffers can be tested in each staining
solution.
STAINING
Specific activity stains are used to detect the
location of specific enzymes after completion of
electrophoresis, The detection of the activity of
individual enzymes is possible because the appropriate
substrates and cofactors required for activity are
provided in the staining solution; the enzymatic reaction
then forms a colored product, either through direct
activity on a dye or by involving other enzymes in a
series of reactions with the generation of a colored
product as the final result. Fluorescent products can be
detected with ultraviolet light; alternatively, non-
fluorescent products can be visualized as a negative
stain by reacting the starch with a ée fluorescent
compound. The biochemistry of such staining reactions is
discussed in Gabriel (13).
Staining recipes for 43 commonly studied enzymes are
presented in Appendix III; additional staining recipes
can be found in Harris and Hopkinson (15), Cheliak and
Pitel (7), Conkle et al. (10), Gabriel (13), and Shaw and
Prasad (35). We weigh stain components into pre-labelled
Styrofoam cups during the electrophoretic run. This
process can be done quite rapidly if stains with similar
reagents are grouped together. The amounts of certain
inexpensive reagents can be estimated; precise
measurements are usually not necessary since excess
reagent will not affect the reaction. One or two key
enzymes or cofactors should be left out initially to
prevent premature reactions; these then can be added just
before the staining solution is poured into the staining
tray.
Once the stain has been poured into the staining
tray, the tray should be lightly shaken so that the gel
418
419
slice can move freely in the staining solution, thus
allowing maximum contact of the staining solution with
both surfaces of the gel. The trays then are placed in a
37° C oven to speed development of the enzyme bands.
Since many stains are sensitive to light, the oven door
should be kept closed as much as possible. Agar overlays
are frequently used for enzymes of low activity or low
concentration; this procedure is described in Appendix
III.
GENETIC INTERPRETATION
Genetic interpretations of electrophoretic banding
patterns can produce large amounts of information about
the nuclear condition and genetic make-up of the
organism. A brief summary of the principles of genetic
interpretation is provided (Figs. 3-5); the reader also
is referred to more detailed accounts (12, 15).
Isozymes can be defined as multiple molecular forms
of a given enzyme; these forms usually have similar, if
not identical, enzymatic properties. Isozymes can occur
within a single individual or among different individuals
of the same species. MKIsozymes also can be localized in
specific tissues of an organism or be compartmentalized
within different areas of the cell. Only those isozymes
with amino acid compositions of different net charge, or
those which result in large differences in the shape of
the enzyme, will be differentiated by electrophoresis.
This represents only one third of all possible isozymes
which may be present within a genetic system.
Detectable isozymes can arise from three different
genetic and biochemical phenomena: 1) multiple allelism
at a single locus; 2) multiple loci coding for a single
enzyme; and, 3) post-translational processing and the
formation of secondary isozymes (15). All of these
situations must be considered when trying to interpret
Fig. 2. A) Analytical gel following removal from mold.
B) Gel sliced to make 8-10 thin layers, each to be
stained for a separate enzyme. C) Gel layer to be
stained on a glass plate using an agar overlay. D) Gel
layer during staining in staining tray. E) Gel layer
following staining for mannitol dehydrogenase to detect 2
alleles and E) Six gel pieces stained for glucose-6-
phosphate dehydrogenase in a gel-buffer screen.
420
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421
electrophoretic data.
1. Multiple allelism at a single locus: Variations
in banding patterns frequently are caused by’ the
expression of multiple alleles at a single locus, each
allele coding for a _ structurally distinct polypeptide
chain. The actual banding pattern is dependent on the
number of alleles present at the locus, which in turn is
dependent oon the nuclear condition (monokaryotic,
dikaryotic), ploidy number (haploid, diploid, polyploid),
and genetic make-up (homozygous, heterozygous) of the
organism. Individuals which are haploid or homozygous
will produce simple banding patterns due to’ the
expression of a single allele. Organisms which are
diploid and heterozygous will produce more complex
banding patterns due to the expression of two separate
alleles.
The banding pattern is also dependent on _ the
quaternary structure of the enzyme. Enzymes which are
monomeric are composed of a single polypetide chain;
monomeric isozymes of a heterozygote will appear as a
mixture of the isozymes produced by the two corresponding
homozygotes. Multimeric enzymes consist of two or more
polypeptide chains; heteromeric (hybrid) bands will be
produced by heterozygous individuals as the result of
random combination of polypeptide chains to produce
hybrid enzyme molecules. Heteromeric bands cannot occur
unless different alleles (coding for different poly-
peptide chains) are present in the same individual. The
expected banding patterns for monomeric, dimeric, and
tetrameric enzymes coded by two alleles at a single locus
are presented in Fig. 3. The frequency of occurrence of
the different isozyme molecules, assuming random combi-
nations of polypeptide chains, should follow Mendelian
ratios; the ratio of isomers formed in heterozygotes will
Figure 3: Predicted banding patterns for one locus with
two alleles (A and A"') and two segregating loci which
share the same two alleles for monomeric, dimeric and
tetrameric enzymes of a diploid organism. Genotypes are
listed beneath each banding pattern (phenotype); alleles
are designated by capital letters. The subunit
composition of each protein band is shown on the right;
lower case letters refer to subunit designations. The
expected ratios of banding intensity for each phenotype
is presented beneath the genotype. Adapted from May (19).
422
be 1:1 for monomers, 1:2:1 for dimers, 1:3:3:1 for
trimers and 1:4:6:4:1 for tetramers. In some cases, a
particular polypeptide will not contribute equally to the
activity of the enzyme due to a slow rate of synthesis,
low stability, or a tendency to break down before it can
assemble into the final enzyme. A polypeptide can also
reduce the effectiveness of the enzyme by decreasing its
stability or reducing its catalytic ability; such enzymes
would not be detected in the expected ratios of intensity
on the electrophoretic gel. A complex series of banding
patterns is produced when three or more alleles are
present in a population of organisms (15). Further
examples are demonstrated in Fig. 4.
2. Multiple loci: Multiple loci can also code for a
structurally distinct series of isozymes; up to four
different loci have been reported for some enzymes.
Multiple loci are thought to originate from gene dupli-
cations and subsequent divergence by point mutations;
they are frequently expressed in different tissues of an
organism or compartmentalized indifferent areas of the
cell. Isozymes coded by different loci are frequently
detected in separate regions of the gel due to their
greater differences in charge and conformation than
usually associated with multiple alleles at a _ single
locus. Heteromeric bands can be formed by polypeptides
coded by different loci; banding patterns thus can become
quite complex when a number of different loci and alleles
are involved (Fig. 5). The distribution of loci is
usually constant within a given species (15).
3. Secondary isozymes: Banding sequences may not
appear to follow expected genetic patterns due _ to
post-translational processing and other events which form
secondary isozymes. Common modifications include _ the
deamidation of glutamine and asparagine residues,
acetylations, oxidation of sulfhydryl groups, additions
and removals of carbohydrate and phosphate moieties,
cleavage of the enzyme by proteases, and the aggregation
or polymerization of the molecule. The formation of such
secondary isozymes is usually quite uniform and can be
recognized by the production of a series of bands for
each allele (15).
Another process responsible for the generation of
secondary isozymes is conformational isomerism. Some
enzymes may have several stable configurations which vary
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423
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GENOTYPE
Fig. 4: Predicted banding patterns of dimeric (A) and
tetrameric (B) enzymes coded by a single locus and three
electrophoretically distinct alleles (A, A' and A'') of a
diploid organism. Genotypes are listed below each
banding pattern (phenotype). The subunit composition of
each protein band is shown on the right; lower case
letters refer to subunit designations. Adapted from May
CLI).
in tertiary or quaternary structure; these will
frequently have different electrophoretic mobilities.
All configurations should be isolated from a single
preparation, and a series of electrophoretic bands should
be observed for a single allele (15).
Enzymes which require cofactors, such as NAD and
NADP, may vary in their electrophoretic mobility
depending on the degree of saturation of the enzyme with
424
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COMPOSITION
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GENOTYPE
Fig. 5: Predicted banding patterns of a dimeric enzyme
coded by two loci which share the same three electro-
phoretically distinct alleles (A, A' and A''). Genotypes
are listed below each banding pattern (phenotype). The
subunit composition of each protein band is shown on the
right; lower case letters refer to subunit designations.
Adapted from May (19).
the cofactor. The amount of cofactor in the staining
solution must not be limiting or inconsistent results are
likely to occur. It may be necessary to add the cofactor
to the sample buffer and/or the gel buffer in order to
maintain the stability of the enzyme during electro-
phoresis (15).
Artifact bands can also arise from the proteolysis of
the sample during extraction and storage; sample material
can be preserved at -80° C or below, however samples
should not be kept for more than one year as activity
will generally decrease. Enzymes can also be bound to
other cell constituents and will not migrate freely in
the electrophoretic field. Alternate methods of sample
extraction and storage may be necessary; stabilizing
agents may be added to the sample to protect the enzymes
from other cell constituents (15).
425
GENIC NOMENCLATURE AND PRESENTATION OF RESULTS
Taxonomic and genetic information can be summarized
by several different methods; the technique of May et al.
(23), as further described by Bonde et al. (3) allows
direct comparisons from one experiment to another
regardless of absolute migration distances if standard
alleles are included in each test. Alleles at a single
locus are described by their relative anodal or cathodal
movement from the origin as compared to the movement of
the homomeric protein coded by the most frequently
observed allele (which is designated 100). A dimeric
heterozygote (or heterokaryon), which produces’ three
bands on the electrophoretic gel, would be designated by
two different alleles (e.g. 100/130); a diploid
homozygous individual would be described by two identical
alleles (100/100 or 130/130). The frequency of shared
alleles can be used to calculate the coefficient of
similarity among organisms. Several different formulas
can be used to determine this quantity (12, 30);
commercial computer programs are also available to do
these calculations. The final information can _ be
summarized in the form of a dendrogram; this type of
diagram can be constructed with a variety of clustering
procedures (11, 12) using commercial computer programs.
TROUBLESHOOTING
Problems sometimes develop during electrophoresis
that threaten the success of the run or result in
inconsistencies among runs performed at different times.
These problems frequently can be corrected or avoided if
proper steps are taken. A list of common problems, along
with their probable causes and possible corrections, is
presented below.
I. Problems with preparation of starch solution.
A. Flecks or lumps appear in hot solution.
1. Cause:
a. Starch was not mixed well with cold buffer.
b. Starch was allowed to settle in the cold
buffer after it was mixed.
c. Heated buffer was not hot enough when poured
into cold starch suspension.
426
d. Flask was not thoroughly cleaned from
previous experiment and flecks of dried
starch have come off the glass.
2. Correction: Discard solution and begin again,
making certain that the starch is mixed well in the cold
buffer, that the remaining buffer is brought to a rolling
boil, and that all glassware is thoroughly clean.
B. Starch is thick and lumpy when poured into gel
mold.
1. Cause: Starch has cooled during the degassing
process.
2. Correction: Discard the solution and repeat
gel preparation. Degas solution for shorter period of
time.
II. Problems with gel consistencies or dimensions.
A. Gels soft and watery.
1. Causes:
a. Starch not cooked long enough.
b. Improper amount of gel buffer added to
starch.
c. Starch not degassed long enough.
d. Gels are still too warm and need additional
time to cool.
e. Gels allowed to warm up during electro-
phoresis process.
2. Corrections: Repeat gel preparation, making
certain that the solution is mixed properly. Increase
periods of heating and degassing; be certain that the
gels are cooled sufficiently before they are trimmed.
Make certain that cooling apparatus is adequate for
electrophoretic conditions. It may be necessary to
decrease current and voltage and allow the gel to run for
a longer period of time.
B. Gels hard and difficult to slice.
1. Causes:
a. Starch cooked or degassed too long.
b. Gel frozen in center from ice packs.
2. Corrections: Determine whether center of gel
is frozen; uneven electrophoretic mobilities can _ be
expected and comparisons will be difficult to make in
this event. Avoid this in future runs by placing glass
plates beneath ice pack or otherwise altering cooling
apparatus. If gel is not frozen, repeat gel preparation
427
while decreasing amount of heating and degassing.
C. Gel uneven in thickness.
1. Causes:
a. Gel mold is not level.
b. Plexiglass strips are not firmly clamped
forming a slow leak on one side of the gel.
2. Correction: Repeat gel preparation making
certain that gel mold is level and that clamps are firmly
in place.
III. Problems with establishing proper voltage and
current.
A. No current or voltage registered on power supply.
1. Causes:
a. Power supply not plugged in, with blown
fuse, or malfunctioning.
b. Broken wire in anode or cathode lead.
2. Corrections: Attach gel to second power sup-
ply with the same leads; if voltage and current still do
not register, repeat process with another set of leads.
B. Current reading is zero at proper voltage.
1. Cause: Corrosion on electrode prevents contact
with buffer.
2. Correction: Remove corrosion with file; re-
solder platinum wire if necessary.
C. Current reading is infinite, voltage reading is
correct.
1. Causes:
a. Short circuit in power supply, electrode
leads, buffer systems or gel.
b. Incorrect settings on power supply.
2. Corrections: Substitute a second power supply
to determine whether short circuit is in gel or power
supply. With power turned off, look for possible short
circuit in electrode leads, buffer or gel systems. Check
settings on power supply to see if they fall within the
proper range (i.e. set at 0-200 mA instead of 0-20 mA).
D. Voltage is too high (or current is too low).
1. Causes:
a. Separation of gel at origin.
b. Poor contact of absorbant cloths with gel
surface.
428
c. Obstruction in flow of electrode buffer
through absorbant cloths.
d. Electrode buffer at improper ionic strength
- too low.
e. Not enough electrode buffer in tray.
2. Corrections: After turning power off, remove
plastic wrap and realign absorbant cloths on gel surface
making certain that there are no wrinkles and folds which
would restrict flow of electrode buffer. Recover with
plastic and turn power on. If voltage is still too high,
add more electrode buffer to tray. If this also does not
correct the problem, run the gel as it is but allow it to
go for a longer period of time.
E. Voltage is too low (or current is too high).
1. Causes:
a. Electrode buffer at improper ionic strength
- too high.
b. Too much electrode buffer in tray.
2. Corrections: Remove some of electrode buffer
from tray. If this does not affect voltage, run gel for
shorter period of time. Be certain that gel does not
overheat; do not allow current to go higher than that
described for the particular buffer.
IV. Problems with uneven migration of dye front.
A. Dye front is concave when viewed from origin.
1, Cause: Uneven cooling of gel; center of gel
is cooler than edges.
2. Correction: Modify cooling system so that gel
temperature does not exceed 4° C and gel is cooled
uniformly.
B. Dye front is convex when viewed from origin.
1. Cause: Uneven cooling of gel; center of gel
is warmer than edges.
2. Correction: Modify cooling system so that gel
temperature does not exceed 4° C and gel is cooled
uniformly.
C. Dye front is straight but higher on one side than
the other.
1. Cause:
a. Gel of unequal thickness.
b. Anodal electrode is worn or corroded.
c. Gel has separated at origin.
429
d. Plastic wrap is not positioned in straight
line along the edge of the gel.
2. Correction: After power is turned off, remove
plastic wrap and check whether the gel has separated at
the origin. Realign plastic wrap and continue run.
Replace worn and corroded electrodes.
D. Dye front is wavy and uneven.
1. Cause:
a. Uneven cooling of gel.
b. Gel has separated at origin.
c. Absorbant cloths not making proper contact
with gel.
2. Correction: After power is turned off, remove
plastic wrap and check whether gel has separated at
origin. Reposition absorbant cloths, making certain that
they are in proper contact with the gel. Modify cooling
system so that gel temperature does not exceed 4° C and
gel is cooled uniformly.
E. Dye front migrates slower in one gel than another
gel of the same systen.
1. Cause:
a. Gels are of different thicknesses.
b. Electrodes are corroded.
c. Unequal levels of electrode buffers are
present in the trays.
d. New absorbant cloths are being used.
2. Correction: Decrease voltage of faster run-
ning gel or allow the slower gel to run for a longer
period of time.
F., Dye front migrates into narrow, “cathodal” sec-
tion of the gel.
1. Cause: Electrode leads have been reversed in
buffer wells.
2. Correction: Reverse leads and allow gel to
run for an additional period of time if this is
discovered early in the electrophoretic process. Not
much can be done if this reverse is discovered towards
the end of the procedure; the "cathodal" strip can be
stained but enzyme separation is usually not reliable.
V. Problems with resolution of protein bands.
A. Bands smeared or streaked from origin.
1. Cause:
430
a. Insoluble fungal material on wick or cut
surface of origin continuously supplies
protein to gel.
b. Low ionic strength of sample buffer allows
precipitation of proteins in sample.
Precipitation will also occur if not enough
sample buffer is added to the crude
homogenate.
c. Air bubble in gel, or gels have separated
at origin.
d. Amylase may be present in crude homogenate.
2. Correction: Since streaking is not detected
until after the gels have been stained, there is nothing
that can be done to save the preparation. In future
electrophoretic runs, be certain that wicks do not
contain insoluble material and the proper amount of
sample buffer is added. Examine gels for air bubbles
while starch is still molten and make certain that the
gels meet firmly at the origin.
B. Bands with poor resolution and poor separation.
1. Cause:
a. Gel and electrode buffers at improper pH or
fonic strength.
b. Wicks are too large so that too much sample
is applied to the gel.
c. Samples permitted to diffuse before or
after electrophoresis.
d. Gel developed too long in staining solution.
2. Correction: Conduct a screening run to deter-
mine which buffer systems should be used with a partic—
ular enzyme. Do not use oversized wicks. Do not allow
gels to sit for long periods of time before or after
electrophoresis. Analyze gels rapidly; photograph each
gel as soon as it is developed.
C. Protein bands very light and difficult to see.
1. Cause:
a. Sample too dilute either through. the
addition of too much _ sample buffer or
improper sample preparation and extraction
procedures.
b. Gel not completely developed in staining
solution.
2. Correction: Allow gels to develop for longer
period of time in the staining solution. If this does
not help, reevaluate procedures of sample preparation.
431
Weak reactions can sometimes be stabilized if the enzyme
substrate, or protease inhibitors, are added to the
sample buffer. The use of agar overlays may also sharpen
staining reactions.
D. Protein bands do not move away from origin.
1. Cause: Improper pH or ionic strength of
electrode or gel buffer.
2. Correction: Conduct a screening run_ to
determine which buffer systems should be used for a
particular enzyme. Make certain that buffers have been
made properly.
E. No bands are detected.
1. Cause:
a. Staining solutions have been mixed
improperly.
b. Chemicals or stain buffers have
deteriorated.
2. Correction: Check stain recipes to determine
whether all components have been added; chemicals which
have been left out can often be added later with no
detrimental effects. If this does not work, remix stain
buffers or order new chemicals. Be certain’ that
chemicals are stored under proper conditions.
ACKNOWLEDGMENTS
The authors wish to express their appreciation to
photographer Austin D. Twigg, TASC, Fort Detrick and
secretary Gwen Bowlus, USDA, FD-WSRU, for their excellent
contributions toward the preparation of this manuscript.
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Appendix II: Buffer systems commonly used in
electrophoresis.
Many different electrophoretic buffer systems have
been published; only a few are presented here. Buffers
routinely used in the Foreign Disease-Weed Science
Research Unit are indicated. Formulas for additional
buffer systems can be found in: Selander et al. (33),
Werth (41), Shaw and Prasad (35), Cheliak and Pitel (7),
Siciliano and Shaw (37), and Conkle et al. (10).
Sample buffer for protein extraction - 0.05 M Tris-Cl
buffer, pH = 7.1.
Ridgway (29) buffer, pH 8.5/8.1:
Gel stock solution, pH 8.5 Electrode buffer, pH 8.1
0.030 M Tris 0.06 M Lithium
0.005 M Citric acid hydroxide
0.30 M Boric acid
Gel buffer made with 99% gel stock solution and 12%
electrode buffer. Run at 250 V; 75 mA
Clayton and Tretiak (8) morpholine buffer, pH 6.1
Gel buffer Electrode buffer
Dilute electrode buffer 0.04 M Citric acid,
1:10 titrated to pH 6.1
with N-3(3-amino-
propyl)-morpholine.
Run at 200 V; 90 mA
Cheliak (7) buffer, pH 7.0/7.0
Gel buffer, pH 7.0 Electrode buffer, pH 7.0
0.05 M histidine-HCl 0.125 M Tris
0.0014 EDTA (pH 7.0 with 1.0 M
(pH 7.0 with 1.0 M citric acid)
Tris)
Run at 50 mA; 3 hrs
439
Steiner (39) buffer, pH 8.4/8.1
Gel buffer, pH 8.4 Electrode buffer, pH 8.1
0.074 M Tris Leo aMelris
0.009 M citric acid 0.314 M citric acid
Dilute cathode tray 1:3
Dilute anode tray 1:4
Run at 90 mA - 3 hrs
May's (19) modification of Selander's buffer, pH 6.3/6.7
Gel buffer, pH 6.7 Electrode buffer, pH 6.3
0.006 M Tris 0.068 M Tris
0.003 M Citric acid 0.037 M Citric acid
Run at 250 V; 75 mA.
May's (19) modification of Markert and Faulhaber's buffer
(18), pH 8.7
Gel buffer Electrode buffer, pH 8.7
Dilute electrode buffer 0.180 M Tris
1:4 0.100 M Boric acid
0.004 M Disodium EDTA
Run at 275 V; 70 mA
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MYCOTAXON
Vol. XXVII, pp. 451-497 October-December 1986
THE LICHEN FLORA OF THE GALAPAGOS ISLANDS, ECUADOR
WILLIAM A. WEBER
University of Colorado Museum
Campus Box 218, Boulder, CO 80309, USA
SUMMARY
196 species of lichens are reported for the Galapagos
Islands, of which eleven, or five per cent, are believed to be
endemic. One new species, Stereocaulon azulense Yoshimura &
Weber, a close relative of the Beringian Arctic S. wrightii
Tuck. and S. apocalyptum Nyl., is proposed, and one new
combination, Buellia mamillana, is made for B. glaziouana.
The Galapagos Islands need no introduction. The visit of
Charles Darwin and the H.M.S. Beagle in 1835 was integral to the
development of modern understanding of speciation and evolution,
and Galapagos is its showcase. Countless tourists know the area
first-hand through the development of the National Park system
of Ecuador and the Charles Darwin Research Station on Isla Santa
Cruz. Many popular and semipopular books have been written
about the flora and fauna, and television has brought the
islands into every home and schoolroom.
A vascular flora of the islands (Wiggins & Porter 1971)
provided a modern base for research which has stimulated many
continuing studies of taxonomy and ecology. Gradstein & Weber
(1982) discussed the plant geography of the bryophytes, and
Weber (1976) published an updated catalog of the mosses. Intro-
ductory outlines of the lichen flora have been provided for
general readers (Weber, Gradstein et al 1977) and for Spanish-
speaking students (Weber & Lanier 1977). Weber & Beck (1986)
discuss the effects on cryptogamic vegetation of the 1982-1983
El Nino phenomenon.
Contribution No. 397 of the Charles Darwin Foundation
452
An excellent description of the islands, their vegetation,
and the history of botanical exploration, including all of the
expeditions upon which lichens were collected, is available in
Wiggins & Porter (1971, pp. 1-45) and need not be repeated here.
The most recent comprehensive taxonomic account of the li-
chens of the Galapagos Islands was Weber (1966). Because of an
imposed editorial constraint, it could include only a historical
list of lichens reported from the islands before the Galapagos
International Scientific Expedition of 1964 took place. All
published records were accounted for, but not all of the speci-
mens of record had been examined for verification purposes.
Since that time, a short paper describing novelties (Weber 1971)
and the publication of records through issuance of the Lichenes
Exsiccati COLO, summarized by Weber (1981), have augmented the
original list. Popular accounts of the lichens were published
also (Weber et al., 1977, 1982).
I have made six collecting trips to the islands, and while
many species remain unidentified, it is desirable to bring our
knowledge of the lichen flora up to date. A number of signifi-
cant reports are contributed here based on the collections on
Isla Santiago by Larry Pike during expeditions sponsored by the
University of Oregon. Incidental collections were made by
various scientists visiting islands during the GISP expedition.
These include Yale Dawson, David Cavagnaro, and Syuzo Itow.
During my subsequent expeditions, I have been aided in the field
by my students, Hans Beck, Jeannine Lanier, Claudia Rector and
Lynn Riedel. The expedition of 1975 was a joint bryological-
lichenological study with my colleagues S. Rob Gradstein and
Harrie Sipman, University of Utrecht, supported by a grant from
the National Geographic Society (Weber 1984a). I am indebted to
my friend Tui De Roy Moore for her occasional and often vital
help in locating rare species.
Santa Cruz Island is probably the richest area for lichen
observations because of its well-developed vegetational zonation
and its accessibility. Coastal rocks near Academy Bay are abun-
dantly covered by lichens. These are best developed where the
rocks are sloping or vertical so as to escape constant heating
and to intercept fog. Coastal shrubs support large populations
of Ramalina and Roccella babingtonii, while mangroves support a
few colorful crustose types. Smooth-barked trees such as Bur-
sera, Pisonia and Piscidia, as well as the trunks of Jasmino-
cereus and the pads of Opuntia, provide excellent lichen sub-
strates. The many cultivated and escaped exotic trees of the
453
cultivated zone, as well as native species of Scalesia, provide
excellent substrates for lichens. Living fences of Jatropha
curcas in the farming area have a wide assortment of lichens,
although most of these shrubs were destroyed during the 1982 El
Nino. The shrubs of the Miconia zone and the summit Scalesia
forests are the best places for large foliose lichens (Sticta
and Pseudocyphellaria). The summit cinder cones support Stere-
ocaulon species. The summit "grassland", dominated in fact by
Cyperaceae, supports clumps of Cladina, Cladia, and many taxa of
"Parmelia" in the broad sense.
Most of my intensive collecting was done within a day’s
walk from the Darwin Station. Other islands on which I myself
have collected lichens are Champion, Espafiola, Fernandina,
Floreana, Isabela, Rabida, Santa Fé, and San Salvador. I am
indebted to Forrest Nelson for his logistical support, and to
the Charles Darwin Station for arranging for fishing boats, and,
in 1984, our circumnavigation of Isabela and Fernandina islands
aboard the Beagle IV, during which my party assisted Carlos
Valle in his census of Galapagos Penguins and Flightless
Cormorants. We appreciated Carlos’ companionship and help to us
during our climb of Cerro Azul.
I also must acknowledge the help, great and small, given us
by the our resident friends who provided food and shelter or led
us to springs and special lichen localities: André and Jacque-
line De Roy and their daughter Tui; Mrs. Horneman and her son
Siegvard; Alf Kastdalen, whose knowledge of the local plants was
unsurpassed (Alf was accidentally electrocuted during a storm in
1982); several children of Carl Angermeyer; and Mrs. Margaret
Wittmer of Floreana. Finally, I appreciated the companionship
in the field of Dr. Ira Wiggins when the two of us decided, in
1964, to work Santa Cruz on foot rather than to gamble and
possibly waste much time jockeying for space and time on side
trips elsewhere.
HISTORY OF COLLECTIONS
Slevin (1959) provides an account of the history of explo-
ration of the Galapagos Islands. While incomplete, his "selec-
ted" bibliography lists most of the pertinent published papers.
If Hugh Cuming or David Douglas collected any lichens on Gala-
pagos, they were probably large fruticose types which were gath-
ered along with vascular plants. Hooker (1846) listed three
lichens collected by Darwin on San Salvador (Santiago or James
Island): Usnea plicata (probably Ramalina usnea, a pendent
lichen which Darwin felt constituted a major element in the food
454
of tortoises), Borrera (=Heterodermia) leucomelos, and Sticta
(=Pseudocyphellaria) aurata. Farlow (1902) contributed a short
lichen list in Robinson’s flora. Some of the earliest, and most
dubious, collections in the record are those made by a Rev. T.
Hill (the chaplain?), on the Agazziz expedition of the Museum of
Comparative Zoology, Harvard University, in 1873, aboard the U.
S. Coast and Geodetic Survey steamer "Hassler". The expedition
spent only nine days anchored at James Bay on Isla Santiago, few
specimens were gathered and collection data were minimal. In
fact, a number of specimens marked "Galapagos" belong to species
endemic to the southern coast of South America. The rock sub-
strates on which they occur are not volcanic, and for practical
purposes most of these should be ignored, except for the fact
that a few new species were based on them. Their type locali-
ties may remain forever unknown.
Previous to the Galapagos International Scientific Expedi-
tion of 1964 (Bowman, ed. 1966), only one lichen student had
visited the island: Albert W. C. T. Herre, who landed in Janu-
ary, 1929 at Post-Office Bay on Floreana and at Academy Bay on
Santa Cruz, making a few casual lichen collections. Being an
amateur lichenologist as well as a professional ichthyologist,
his collections, though few, were out of the ordinary, yielding
a few undescribed species and one notable disjunction. Most
collections had been made casually by scientists of other dis-
ciplines. For example, in 1898, Edmund Heller and Robert E.
Snodgrass, zoologists of Stanford University aboard the schooner
"Julia E. Whalen", collected animals and plants, some of which
were lichens, from December to the following June.
Serious lichenology began on Galdpagos with the expedition
of the California Academy of Sciences, 1905-1906. The botanist
of the expedition was Alban Stewart, an instructor in Botany at
the University of Wisconsin. His lichens were named by W. G.
Farlow and are housed at the Farlow Herbarium, Harvard Univer-
sity. Stewart's own words give some indication of the extent to
which he involved himself with lichens. "I am not a lichenolo-
gist in any sense of the word, and it is with some hesitation
that I approach a subject with which I have so little acquain-
tance. However, as I made a number of notes on the subject
while collecting there, and as very little has been written on
the general distribution etc. of the Lichenes of these islands,
it seems worth while to publish these notes, along with a list
of the species which were secured."
"This list is probably far from complete, for I was more
interested in collecting and studying the distribution of the
455
vascular plants; and it was often the case that lichens were
carefully collected only where no vascular plants in good con-
dition were to be found, as was sometimes the case when the
lower islands were visited during the dry season. Notwith-
standing the rather neglectful way in which this group of plants
was treated, sixteen species were found which had not before
been reported from the islands, and the range of quite a number
of species already known was considerably extended. There were,
however, some fifteen species, reported by former expeditions to
the islands, which do not appear in my collection. Some of
these have been taken but once."
One can read between the lines that, as is often the case,
botany would be low in the peck order on an expedition dominated
by zoologists intent upon collecting Galdpagos land tortoises.
Stewart might have been frustrated very often at being unable to
stay in a rich botanical area when the zoologists had decided
they had completed their work. With his major goals fixed on
the vascular plants, we can only be grateful to him for devoting
even a small portion of his time to collecting lichens.
Svenson (1963) decried the lack of attention given to li-
chens and singled them out as being in great need of study on
the islands. Farlow (1902) had written: "Judged by the
collections of previous expeditions as well as those of Messrs.
Snodgrass and Heller, the lichen flora of the Galdpagos Islands
must be striking in appearance and abundant. Naturally the
collections include principally the larger and more showy
species of foliose and fruticulose habit, which are in most
cases identical with species found along the Pacific Coast of
America from California to Chile." Swenson (1935), describing
Tower Island, says: "The trees...have swollen and twisted
branches cluttered with white coralliform lichens." And Stewart
(1912) wrote: "When one lands for the first time on almost any
of the islands, one is immediately struck with the great
abundance of lichens. This is true not only of the larger and
higher islands, which reach sufficient elevation to receive a
considerable amount of moisture from the fog-banks which strike
their sides...but it is true also...of the lower islands...
where...desert or semidesert conditions prevail."
In the winter of 1963, I was scheduled to make a trip into
Baja California with Reid Moran, but he suggested that I get in
touch with Bob Bowman, who was organizing what became the Gala-
pagos International Scientific Expedition (GISP), a multidisci-
plinary scientific group planning to go to the Galapagos for
several reasons: to dramatize the importance of the Galapagos
456
as a unique scientific area from a biological and historical
viewpoint as the birthplace of the theory of Natural Selection,
to dedicate the new Darwin Research Station there, and through
seminars, to explain to the people of Ecuador what a treasure
they had and the necessity to preserve it for posterity.
When I called Bowman and asked if they had planned to take
along a cryptogamic botanist he retorted, "No, should we?".
Fortunately, he was very easy to convince, and so began the
first visit to the islands by a lichenologist. I shall never
forget my first impressions of the lichen flora. I had gleaned
all of the published literature on the islands and found only a
single illustration showing lichens--a cover of "Life" magazine
showing a leaning manzanillo tree upon which the photographer
had posed a land iguana. The bark of the tree appeared to have
crustose lichens on it.
Upon landing at Puerta Ayora in Academy Bay I was detailed
for a few hours to stand guard over the luggage, but at the
first opportunity I walked a few hundred yards back of the Dar-
win Station into the cactus forest, which at that time did not
have neat crushed coral walks and man-made walls. I was quite
overwhelmed. Every rock, cactus pad and tree trunk seemed to be
to be totally covered with white and varicolored foliose and
crustose lichens, and I sat down and almost wept because I felt
that I couldn't possibly get on top of this fantastic lichen
flora in the six weeks at my disposal. Bob Usinger, in his
autobiography (1972) came away with a slightly different im-
pression of my feelings, writing: "[Weber] came in from his
first day of field work like a small boy from a candy shop."
I feel very fortunate that, since 1964, I have been able to
revisit the islands five times, sometimes leading student tours,
and sometimes on collecting trips. I have been able to collect
the majority of the obvious species and some of the cryptic
ones. Nevertheless, as in any area, the real floristic picture
will only emerge when a well-trained and acutely perceptive
resident lichenologist spends extended periods on all the
islands, and when enough monographic work has been accomplished
to make possible the identification of the difficult groups. I
sincerely hope that this will be accomplished through an
Ecuadorean botanist-in-residence.
A general description of the lichen vegetation is not given
here because I have developed it elsewhere (Weber & Gradstein,
ibid.). Some observations should be made concerning endemism in
general and inter-island endemism. In the vascular plants,
457
inter-island endemism is quite important. In the lichens I be-
lieve that a given lichen species will be present on any island
which meets its ecological requirements, and that there is no
evidence that natural selection has operated to produce special
island races or species.
Significant Galapagos endemism occurs in the lichen flora,
nevertheless. This is most apparent in the crustose and fruti-
cose lichens inhabiting the coastal rocks. There are endemic
species, often with close relatives on the coast of Chile, and
there are a few disjunct species involving Galapagos and main-
land patterns. The lichens growing on the bark of trees tend to
belong to more widely distributed Neotropical species, and only
a few taxa in the highest reaches of the largest islands are
Andean. I have developed this discussion elsewhere (Weber &
Gradstein, ibid.).
I did not realize until my expedition of 1983 (Weber & Beck
1986) what a devastating effect the extremely high rainfall of
the record El Nifio event might have on lichens, when running
fresh water flowed over the coastal rocks for many days at a
time, leaching pigments and scouring the cortices of crustose
lichens so that they became indistinguishable, waterlogging the
foliose and fruticose lichens so that they fell from the trees
in masses, and saturating the ground and the bark of trees, re-
sulting in catastrophic losses of habitats.
In this connection, Hale (1984), based on studies of the
flooding of a fresh-water stream in Florida, found that some
corticolous lichens are killed by submergence of a few days.
Santesson (1939) suggested that one to two weeks of submergence
was a likely minimum for lacustrine saxicolous lichens in
Sweden. During the Great El Nifio there was a combination of
submergence in salt water caused by prolonged abnormal high
tides and prolonged fresh-water runoff. Even on elevated ground
the lichen cover of rocks was leached and decorticated.
Hamann (1986) noted a depression in a lava field on Santa
Fé Island several years before the Great Nifio which was bordered
by a prominent upper line of saxicolous lichens. It occurred to
him that at some previous period this depression must have been
a lake below the highest level of which all lichen growth had
been destroyed. His assumption was correct, for the lake was
refilled in 1983!
The ancient high water line of the pluvial period (about
11,000 years ago) in the Great Basin of the western United
458
States at Lake Lahontan, Nevada, is marked by a prominent high
water lichen line of Acarospora chlorophana. Recovery of the
lichen cover below this level is insignificant although the
thriving lichens above the ancient water level can be seen from
several miles distant.
I anticipate that the lichen flora of the Galapagos will
eventually recover, but I expect many decades to elapse before
we will see again the luxuriant lichen tapestry that covered the
islands in 1964.
SCOPE OF THE CATALOG
The catalog lists 196 lichen species that are reasonably
correctly identified up to the present moment. Previous reports
have been investigated and the cited specimens, in most instan-
ces, were located and evaluated. Old names have been brought up
to date. New reports for the Galdpagos Islands are signaled by
an asterisk. All specimens cited, except as otherwise noted,
are in Herbarium COLO, Boulder; the cited number in all instan-
ces is that of the COLO lichen accession book. When reports are
few and from one island, citations are given. When numerous
collections were made and the taxon is not a critical one, only
one or a few specimens are cited and only the names of islands
are given. All numbers distributed in Lich. Exs COLO are cited.
Collections examined at other institutions are listed under
collector’s number and herbarium acronym. Complete citations of
species names are given only when the name is not easily avail-
able in Zahlbruckner’s Catalogus Lichenum Universalis.
The name of the lichen is followed by the collection re-
cords, and a brief statement of the habitat, substrate, and
relative abundance. Very brief statements concerning recogni-
tion are given for the more easily recognized species; they are
not intended to be exhaustive, but are only provided to enable
visitors to Galapagos, who cannot obtain collection permits, to
be able to identify some of the more conspicuous species in the
field. Characteristics only visible by compound microscopy are
omitted. Specialists should refer to the original literature
and to the specimens distributed in Lichenes Exsiccati COLO.
A significant part of the collections remains unidentified,
not because they are undescribed species but because the liter-
ature and collections of Neotropical lichens, especially crust-
ose forms, are scattered and the specimens need to be compared
with type specimens in European herbaria or require the atten-
tion of monographers. The families Arthoniaceae and Graphid-
459
aceae, and genera such as Buellia and Bacidia loom large in this
category. Any of these groups will be made freely available on
loan upon request of the monographer.
Appendix I deals with synonyms, misidentifications and
misapplied names found in the earlier published reports.
Appendix II presents new combinations. Appendix III contains
descriptions of new taxa.
ENDEMIC TAXA
Eleven species are now thought to be Galdpagos endemics:
Arthonia platyspilea, Arthothelium galapagense, Bacidia insu-
laris, Buellia galapagona, B. straminea, Lecanora pseudopinguis,,
Pertusaria albinea, Physcia insularis, Roccella galapagoénsis,
Stereocaulon azulense, S. weberi. The four corticolous species,
however, probably will be found on the mainland. In any event,
the endemism is extremely low, constituting only five per cent
or less, of the known flora.
SPECIES LIST
Note: Names of islands conform to the alternatives chosen
by Wiggins & Porter (1971, end-papers map).
*Acarospora schleicheri (Ach.) Mass.
Rabida: 40100; Santa Cruz, 54819. Infrequent or rare on
lava rocks, particularly those with a sandstone-like texture.
Especially well-developed on Rabida. This is a form with small
angular areoles.
*Anthracothecium leucostomum (Ach.) Malme.
P. libricola Fée, Essai, p. 82. 1837. Santa Cruz, Weber
40199, 41111, 40193, 40255, 40258, abundant on living fences of
Jatropha curcas. Most Jatropha fences on Santa Cruz were de-
stroyed during the Great Nifio. San Salvador: Pike 56180 (on
Acacia rorudiana). Evidently closely related or identical to
A. pyrenuloides (Mont.) MUll.-Arg.
Anthracothecium ochraceoflavum (Nyl.) Willey.
Floreana, Genovesa, Fernandina, San Salvador, Santa Cruz,
Santa Fé, Tower, Wenman. A common crustose lichen making cons-
picuous yellow splotches on smooth-barked trees such as Bursera,
Laguncularia, Rhizophora and Hippomane. Restricted to areas at
sea level and close to tidal influence, extending to the tran-
sition zone in some places. The thallus is pale yellow-orange,
with darker rust-orange perithecia (not to be confused with Py-
460
renula cerina, which has a deep red-orange thallus and almost
invisible perithecia).
Arthonia platyspilea Nyl.
"Galapagos", Hassler Expedition (Willey 1890): Gardner:
Snodgrass & Heller (Farlow 1902); San Salvador: Pike 55200,
55222; Santa Cruz: Lich. Exs. COLO 122. Common on Croton, also
found on Bursera. Stroma pale, irregular, "bruised-looking";
thallus IKI+ blue.
Arthonia tumidula (Ach.) Ach. A. cinnabarina (DC.) Wallr. A.
gregaria (Weig.) Koerb.
Pinzon, San Cristébal, Santa Cruz. Very common on
smooth-barked trees at all altitudes. Recognized by the small
round plane grayish or reddish stroma with red Trentepohlia
algae "leaking out" around the margins or sometimes staining the
entire disk or thallus red.
Arthothelium galapagoénse Huneck & Follmann.
Endemic. Plazas, San Salvador, Santa Cruz: Lich. Exs. COLO
113. A dominant crust on rocks just above the high tide mark,
forming large white patches with irregular-shaped bluish-pruin-
ose stromae. A counterpart of the Chilean coastal A. spiloma-
toides (Nyl.) Zahlbr.
*Aspidophyllum fugiens (Fée) R. Sant.
Santa Cruz: on leaves of Coffea, Horneman garden, 40433,
!Vezda.
*Bacidia apiahica (MUll.-Arg.) Zahlbr.
Santa Cruz: on leaves of Miconia robinsoniana, summit heath
zone, 40216, !Vezda.
Bacidia insularis Zahlbr.
Endemic, the type collected on Floreana, Post Office Bay,
on Bursera, by Herre. Bartolomé, Plazas, Santa Cruz. Common on
Laguncularia, Castela, Bursera (smooth-barked trees) on live
bark and dead decorticated wood.
*Bacidia millegrana (Tayl.) Zahlbr.
Pinta, Santa Cruz: Lich. Exs. COLO 121, San Salvador: Com-
mon on Bursera and Piscidia in the arid zone.
*Bacidia rubella (Ehrh.) Mass. B. luteola (Schrad.) Mudd.
Santa Cruz: on Jatropha curcas living fence, Bella Vista,
ALiZ5"
461
*Bacidia subleporina (Nyl.) Zahlbr.
San Cristobal: on Citrus leaves below Tres Palos, Lanier
63764 (in packet of Tapellaria epiphylla). !Vezda.
*Brigantiaea leucoxantha (Spreng.) Sant. & Hafelln.
San Salvador: Xanthoxylum forest with Psychotria understory
along trail from top of James Bay lava flow top the western
peak, 530 msm., Pike ID18-16. !Hafellner.
*Buellia dejungens (Nyl.) Vain.
Santa Cruz: on stone, barranca behind Darwin Station,
40838. Thought to be endemic to the West Indies, according to
Imshaug (1955).
*Buellia galapagona W. A. Weber, The Bryologist 74:188. 1971.
Endemies))San Salvador: Pike) L-55431; Santa Cruz:bich.
Exs. COLO 344 (type), 40360, 40872. A dominant and conspicuous
coastal rock lichen in association with Lecanora pseudopinguis
and Arthothelium galapagoénse. Forming large tan-brown patches
with large black apothecia.
*Buellia lauricassiae (Fée) MUll.-Arg. See Imshaug (1951,
Pesala).
Santa Cruz: on Bursera, cactus forest, 40336.
Buellia mamillana (Tuck.) W. A. Weber, n. comb. See Appendix II.
Darwin, San Salvador, Santa Cruz, Wolf. An abundant pale
yellow or white crustose lichen growing on dry surfaces of hor-
izontal lava blocks in the cactus forest. The thallus is
minutely areolate, effuse, with small black or somewhat pruinose
convex apothecia. Widespread in subtropical America.
*Buellia modesta (Kremp.) MUull.-Arg. See Imshaug (1955a, p.
B97 ic
Fernandina: 40463; Santa Cruz: 40061, 40841, 40859, 41048.
On lava in the dry cactus forest and on coastal rocks. Imshaug
deals with a corticolous plant, but Malme (1927) reports it in
South America equally on rocks and trees.
*Buellia parasema (Ach.) DeNot. See Imshaug (1951, p. 84-92).
San, Salvador? (On Bursera,..250 mos.m., Pike. 56171.
Buellia straminea Farlow in Robinson. See Imshaug (1955, p.
282-283).
Endemic. Bartolomé, Fernandina, Isabela, Pinta, Plazas,
Rabida, San Salvador, Santa Cruz. While not a dominant or
462
conspicuous species, B. straminea is a constant on coastal
rocks. The type material did not portray the variability of the
species as I know it in nature, the yellow thallus being
distinctly radiate-lobed when well-developed on hard lava but
consisting of flecks and irregular lobulate areoles when
pioneering, for example, on soft tufa. Pyxine glebosa Tuck.,
described from Galapagos and discussed by Imshaug (l.c.) can
safely be placed in synonymy.
*Bulbothrix laevigatula (Nyl.) Hale. Parmelia laevigatula Nyl.
San Cristébal: Wreck Bay, Itow 40982; Santa Cruz: 40402.
!Hale. Thallus white, with bulbate marginal cilia, rounded si-
nuses, narrow lobes; surface isidiate, not maculate; cortex K+
yellow, med. K-, KC+ red (lecanoric).
*Byssoloma subdiscordans (Nyl.) P. James.
Floreana: 64132; San Cristébal: Gradstein 63825 p.p.;
Santa Cruz: Sipman 8, Weber 64131, 40247, 40248, 40250, 63479.
! Vezda. On leaves of Eugenia jambos, zone of cultivation,
with Strigula, Porina, Sporopodium, Echinoplaca and Tapellaria.
*Byssoloma tricholomum (Mont.) R. Sant.
San Cristébal: on citrus leaves below Tres Palos, Lanier
L-63764 (in packet with Tapellaria epiphylla). !Vezda.
*Caloplaca byrsonimae (Malme) Zahlbr. Callopisma byrsonimae
Malme sArk, t7bOt, 2U0A( 9) 33 4970)
Santa Cruz: 40205, 40585 (cum apo.), 40386, 40451, 40563.
San Salvador: Pike 56169. The type came from Matto Grosso,
Brasil. This is a common and easily recognized lichem, usually
a sterile, very thin orange-yellow crust covered with minute
clustered cylindric isidia. The black prothallus is usually
evident in good material. Apothecia are very sparse.
*Caloplaca camptidia (Tuck.) Zahlbr.
San Salvador: Pike’ 55221, 55205, 56168: Santa Cruz: 408637,
Usually on trees, but on boulders at Academy Bay, Santa Cruz.
Apothecia dark brown, with paler rim; epithecium K+ violet wash;
spores 10-12x4-5m , polarilocular with wide isthmus.
*Caloplaca cirrochroa (Ach.) Th. Fr.
Champion: 2781; Santa Cruz: 40055. On shore boulders just
above the high tide mark east of Darwin station, on rock faces
of the barranca behind the station. Thallus orange, lobate but
lobes quite short and broad, erupting back from the tips with
saucer-shaped bowls of yellow soredia; lobe tips somewhat thick
ened, often somewhat pruinose.
463
Caloplaca floridana (Tuck.) S. Tucker, Bryologist 82:132. 1979.
Lecanora floridana Tuck.
San Salvador: on Bursera, Pike 56167. The black-fruited
corticolous Caloplaca species in all probability eventually will
be segregated into new genera.
*Caloplaca isidiosa (Vain.) Zahlbr., Cat. Lich. 3:241. 1931.
Placodium isidiosum Vain., Etude Lich. Brésil 18. 1890.
Darwin, Gardner, Genovesa, Pinta, Plazas, San Salvador,
Santa Cruz, Wolf. Described from Brazil, but evidently the same
taxon occurs on the Chilean coast and in South Africa. A common
orange lichen of coastal rocks, characterized by a narrowly
lobate thallus margin, with slender isidia on the older thallus
parts. Previously reported by Dodge as C. elegans, C. muelleri,
C. rugulosa (Nyl.) Zahlbr., and by Stewart (1912) as Placodium
murorum.
*Chiodecton effusum Fée.
Rabida: on Croton, Cavagnaro 40459.
*Chiodecton farinaceum Fée.
Rabida: 40442: San Salvador: Pike 55411; Santa Cruz: 40155,
GOLD /IUAO LSI) ZO424 40445) 40610. On Laguncularia, cacti,
Bursera, coastal scrub to transition zone.
*Chiodecton myrticola Fée.
Rabida: 40506; San Salvador: Pike 55432; Santa Cruz:
40053, 40607. On Piscidia and Bursera in the cactus forest, up
to 300 m.s.m on Radbida.
*Chrysothrix candelaris (L.) Laundon. lLepraria candelaris (L.)
rr.
Floreana: on Acacia rorudiana, trail from Black Beach to
highlands, 63018 p.p. (with Diploicea canescens), Rabida, Wolf
(on stones). A golden yellow sorediate crust forming large
patches on bark and stones.
*Cladia aggregata (Sw.) Nyl.
Santa Cruz: 55403. In the summit grassland at foot of El
Puntudo, forming large discrete rounded mounds 2-3 dm diameter.
Cladonia-like, with tan to brown, varnished surface, richly
dichotomous, with pointed branch tips; thallus with large per-
forations.
Cladina confusa (R. Sant.) Follm. & Ahti, Philippia 4:314.
1981. Cladonia galapagosensis Ahti; C. polia R. Sant.
464
Fernandina, Isabela, San Cristébal, Santa Cruz: Lich. Exs.
COLO 105 (as C. galapagosensis), 106 (as C. polia). Forming
large hemispherical mounts in the summit grassland, usually an-
chored by entanglement of the thallus with low shrubbery. There
are strains with and without usnic acid. Branching is mostly
anisotomic trichotomic. Reactions with K and Pd are negative.
*Cladina sandstedei (des Abbayes) Ahti, Beih. Nova Hedw. 79:40.
1984. See also Ahti (1961).
Pinta: east slope at 490 m.s.m., much drier zone than sum-
mit sites for Cladina, among Polypodium on old lava flows, di-
rectly above Opuntia bushes, Cavagnaro 68108. Formerly consid-
ered to be endemic to the Caribbean area. Thallus gray, with
the terminal ends blackening; branching anisotomic dichotomic;
cortex K+ yellow, Pd+ orange.
*Cladonia balfourii Crombie
Floreana: 62897, 62988; Isabela: 63523; San Salvador: Pike
56163; Santa Cruz: Lich. Exs. COLO 139. !Thomson. Frequent on
lava knolls with ferns and bryophytes, Transition zone. The
Cladonia chlorophaea group is very difficult, especially in the
tropics. C. balfourii is said to differ from C. chlorophaea by
lacking cups or having very irregular ones. By this criterion
our material could as well belong to C. chlorophaea.
Cladonia ceratophylla (Sw.) Spreng.
San Salvador, Santa Cruz. A handsome and easily recognized
species with large, white, thin basal squamules, convolute when
dry, with long sparse white cilia, no cups but rather acicular
podetia. Common on dead wood and other decaying vegetation in
the zone of cultivation and Miconia belt.
*Cladonia didyma (Fée) Vain.
San Salvador: on shrubs, James Bay, Pike 55241, 55426.
Santa Cruz: summit grassland, 40784. !Thomson. Common in the
summit grassland, often in burned spots. Podetia decorticate
and discoloring dark brown; granular sorediate, K+ yellow, Pd+
red; apothecia red.
*Cladonia macilenta Hoffm. ssp. theiophila Asahina
Isabela?) 63514") Pinzon;) 635/21;\ SannCristobal: (63/35 .6 7290.
Santa Cruz: 63343. !Thomson. On the ground in the summit fern-
sedge zone. Podetia slender, tall, branched, ecorticate,
coarsely sorediate, with large central cavity. Thomson says:
"rather tall, but the chemistry of usnic and barbatic acids in
GE and thamnolic in BaOH is quite diagnostic."
465
*Cladonia pityrea (Flk.) Fries
Fernandina, Floreana, San Salvador, Santa Cruz, !Thomson.
One of the common species of the forest floor, transition and
moist zones.
*Cladonia sphacelata Vainio
Santa Cruz: 63045, 63049. Fern-sedge zone, summit between
El Puntudo and Cerro Crocker. !Thomson. Podetia slender,
blackish, K-, Pd- (in the field I noted that this was greener
and more corticate than 63044 which was growing with it (=C.
subsquamosa) .
*Cladonia cf. subcariosa Nyl.
Santa Cruz: summit fern-sedge zone, on saddle between El
Puntudo and Cerro Crocker, 700 m.s.m., 63339. Thomson says:
"although only sterile squamules, I am thinking of C. subcariosa
Nyl. in the sense of Culberson (1969). The chemical contents,
atranorin plus norstictic acid is not a common combination and
with coarse squamules like this I feel fairly confident of the
determination. Of course, if this turns out to be something
with podetia of quite another kind I would have to revise the
answer. Culberson mentions a Cuban specimen so this may be a
tropical member of the C. cariosa group."
*Cladonia subsquamosa (Nyl.) Vain.
Floreana: on Psidium branches between dense moss cushions,
Gradstein 62997; Santa Cruz: El Puntudo, 63044. !Thomson.
Squamules small, delicate, sorediate, the podetia acicular, so-
rediate, little branched (63044 is ecorticate, esorediate with
squamules on the podetia).
*Cladonia subulata (L.) Wigg.
Isabela: Cerro Azul, S slope above Caleta Iguana, among
fern-sedge vegetation, Sipman 63520. !Thomson.
*Cladonia symphoriza Nyl.
Santa Cruz: saddle between E. Puntudo and Cerro Crocker,
63341 (previously known from Colombia. !Thomson. Podetia stout,
squamulose, with terminal clusters of red apothecia. Mixed with
C. sphacelata Vain.
*Coccocarpia palmicola (Spreng.) Arvidsson & Galloway. See
Arvidsson (1982).
Fernandina, Isabela, Santa Cruz, San Salvador. ! Arvids-
son. Most common on trees, from lower Transition to moist zone,
but occasionally on rocks. Thallus lead-gray with rounded lobes
466
and blue-gray-black rhizines forming a brush below; central part
of thallus with numerous erect cylindric isidia. Our material
always sterile.
Coccocarpia erythroxyli (Spreng.) Swinscow & Krog. C. pellita
var. parmelioides and Pannaria molybdaea of earlier reports.
Santa Cruz: 40290; San Salvador: Pike 2718. !Arvidsson.
Larger than C. palmicola and totally lacking isidia, usually
fertile, with large black convex apothecia.
*Collema furfuraceum (Arn.) DuRietz
Santa Cruz: 40207, 54841; San Salvador: 55428. !Degelius.
Common on Bursera at Academy Bay, near "The Cracks". Thallus
foliose, black, with rounded thallus lobes, ridged-pustulate,
with isidia on the ridges, but the thallus not obscured below
the ridges. Always sterile.
Dictyonema montanum (Sw.) Parmasto ex Follmann. Cora pavonia
(Sw.) Fr.
Santa Cruz. Common on naked or mossy trunks of trees and
shrubs in the moist zone. A conspicuous foliose lichen resem-
bling very thin and fragile shelf-fungi, with surface concentric
arcs, the thalli often imbricate, white when dry, gray-green
when moist. Hymenium yellowish-waxy, on the underside, of dis-
continuous plates or fragments.
Dictyonema guadelupense (Rabenh.) Zahlbr. Dictyonema sericeum
of Howell (1942).
Isabela, San Cristé6bal, San Salvador, Santa Cruz. Infre-
quent on trees in the moist zone, on Miconia robinsoniana.
Thallus dark green, amorphous, with a "nap" of shaggy tan or
whitish hyphal masses.
*Dimerella lutea (Dicks.) Trev.
Santa Cruz: 5 mi E of Horneman Farm, on Psidium galapa-
geium, 40480.
*Diploicea canescens (Dicks.) Mass. Buellia canescens deNot.
Floreana: on Acacia rorudiana bark, Black Beach trail to
highlands, 63018 p.p. (in packet of Chrysothrix candelaris).
*Diploschistes actinostomus (Pers.) Zahlbr.
Santa Cruz: 40043, 40105, 40818, 40825, 40826, 40870. In
frequent, on smooth, hard lava rocks along the shore and on
talus blocks of the barranco near the Darwin Station.
467
*Diploschistes muscorum (Scop.) R. Sant. D. bryophilus (Ehrh. )
Zahlbr.
Isabela: on Stereocaulon azulense, Cerro Azul, 81068.
*Diploschistes scruposus (Schreb.) Norm.
Isabela, San Cristébal, San Salvador. On soil and rocks.
*Dirinaria applanata (Fée) Awasthi (previous reports of Physcia
picta apply to this species).
Genovesa, Radbida, San Salvador, Santa Cruz. Abundant on
twigs and trunks of smooth-barked trees. Thallus closely adnate
with flat, flabellate, confluent and often longitudinally pli-
cate lobes; soralia laminal, round; medulla white. Usually
sterile.
Dirina approximata Zahlbr. Dirina herrei Zahlbr. See Tehlér
C53).
Floreana; Herre in Zahlbr., Lich. Rar. Exs. 269. (Type of
Desnerre:), Lich. Exs, COLO 36/7 (asiD.. herre?): Isabela, Pinta,
Plazas, Rabida, San Salvador, South Seymour: Herre, Krypt. Exs.
Vindob. 3144 (Type). Abundant on Bursera trunks, cacti and
shrubs, in the cactus forest; rarely on rock. This is not an
endemic; we have collections from the adjacent mainland and from
the West Indies.
*Dirinaria caesiopicta (Nyl.) Awasthi, Bibl. Lich. 2:94. 1975.
Bartolomé, Bindloe, Floreana, Genovesa, Isabela, Plazas,
(cited by Awasthi), San Salvador, Santa Cruz. Very common on
coastal rocks.
*Dirinaria confluens (Fr.) Awasthi, Bibl. Lich. 2:28-31. 1975.
(Reports of D. (Physcia) aegialita apply here).
Floreana: 40604; San Salvador: Pike 56160; Without local
ity: Hassler Exped. (FH, cited by Awasthi). Common on bark and
twigs of Bursera. Thallus esorediate, with confluent and pli-
cate lobes; hymenium 80-100m high (the critical difference be-
tween this and D. subconfluens).
*Dirinaria confusa Awasthi, Bibl. Lich. 2:56-60. 1975.
Floreana, Taylor 867 (FH, cited by Awasthi); Genovesa,
Stewart 356 (FH, cited by Awasthi); Rdbida: Stewart 357 [cited
by Thomson (1963) as Physcia aspera]. "Easily distinguished by
the characteristic irregularly thickened, simple to coralloid-
branched crateriform sorediate isidia producing coarse granular
soredia" (Awasthi, 1975). Very similar to D. confluens but
having a lentiform hypothecium, lacking divaricatic acid and
468
said to have the thallus not adglutinated to the substrate.
*Dirinaria leopoldii (Stein) Awasthi, Bibl. Lich. 2:89-91.
Espanfiola: Itow 40736, 40762. On branches of bushes. Si-
milar to D. applanata but with red medulla.
*Endocarpon pusillum Hedw.
San Salvador: on rock, above James Bay, 895 m.s.m., Pike
wo Webs fro wh
*Erioderma sorediatum Galloway & Joergensen, Lichenologist 7:
LI9e Tea Log
Fernandina: 2,000 ft. alt., Snodgrass & Heller (DS), a
fragment 1 x 2 mm found in a packet with Pseudocyphellaria aur-
ata; specimen has a portion of thallus with one apothecium; San
ta Cruz: along trail from Bella Vista to Media Luna, in upper
Miconia zone; on Miconia branchlets in opening near rivulet, 550
m.s.m., Sipman L-21. This is an example of the likelihood of
continual discovery of isolated disjunct species with extremely
small populations in the Galapagos. First described from New
Zealand and reported from the Philippines (Galloway & Joergen-
sen, l.c.), it is known now from Brazil (K. Kalb, Lichenes Neo
tropici 321), and Oregon (Lane Co.: on moss from on Vaccinium
membranaceum in sand dunes, 30 ft. alt., behind Heceta Lodge,
Florence, Howe s n., COLO L-64494). The thallus has bluish
limbiform soralia on the upturned lobe edges; underside white
with bundles of black rhizines.
*Everniastrum vexans Hale, Mycotaxon 3:350. 1976. (Parmelia
americana, P. camtschadalis of reports).
Isabela: Lich. Exs. COLO 499, !Hale; San Salvador, Santa
Cruz. Locally abundant on twigs in humid areas in the high-
lands. Thalli elongate, involute, sparsely black-ciliate, white
above, black and naked beneath.
*Glyphis cicatricosa (Ach.) Zahlbr.
Floreana, Pinzén, San Salvador, Santa Cruz. A thin tan
crust common on smooth bark (Citrus, Scalesia, Croton) in the
moist zone. Very easily recognized. The fruiting area is dis-
crete, almost circular, with many small closely fitting round or
slightly lirelliform brown open disks.
*Graphina virginea (Eschw. in Mart.) MUll.-Arg.
Santa Cruz: Lich. Exs. COLO 646, 40024, Herre 41160, Itow
40733. Forming large continuous white areas on bark of large
avocado tree along fence line just above old Horneman property.
Apothecia elongate, white, not raised.
469
*Graphis afzelii Ach. For Graphis species, see Wirth & Hale,
COMLELO OULU pss NAc Hero LiisO. Ole LOGS TEL es. Gol 46. VWirth
has declined to study the Galapagos material because of pressing
teaching obligations.
San Salvador: on Bursera, Pike 56483.
*Graphis caesiella Vain.
Santa Cruz: Lich. Exs. COLO 480, Santa Fé. A very common
species on Croton but also on Bursera and other trees.
*Graphis desquamescens Fée.
Santa Cruz: on Bursera, Academy Bay, 54823.
*Graphis striatula (Ach.) Spreng.
Santa Cruz: On Castela galapageia, Lich. Exs. COLO 142.
*Graphis subamylacea A. Zahlbr.
Santa Fé: on Opuntia echios, 54830; Santa Cruz: on Bur-
sera, Academy Bay, 54822.
*Gyalectidium filicinum Mull.-Arg.
Floreana: 63344; Santa Cruz: 40211 (with Tapellaria nana),
40216, 40217, 40432, 40433 (in packet of Aspidophyllum fugiens) ,
40434, 63497. !Vezda. On leaves of Miconia, Coffea, and ferns,
in the moist zone.
*Haematomma puniceum (Sm. ex Ach.) Mass. (Lecanora punicea of
previous reports).
Rabida, Floreana, Santa Cruz, San Salvador. Common on Bur-
sera and other trees in the cactus forest zone. An unmistake-
able lichen with white thallus and bright red apothecia.
*Herpothallon sanguineum (Sw.) Tobler. Chiodecton sanguineum
(Sw.))) Vain.
Fernandina, Pinta, Rabida, San Salvador, Santa Cruz. A
unique lichen common on trees of the moist zone, forming great
gray and blood-red splotches.
*Heterodermia albicans (Pers.) Swinscow & Krog, Lichenologist
Cc Loy oO Ae enysCiaralpicans (Pers. ) J. WwW. Thomson:
San Salvador, Santa Cruz, Wolf. Common on living fences of
Jatropha curcas, moist zone. Thalli sterile, white, radiate,
flat or concave, the older parts with marginal soralia; cortex
K+ yellow or yellowish (atranorine & norstictic).
*Heterodermia antillarum (Vain.) Swinscow & Krog, Lichenologist
470
8:114. 1976. Anaptychia tropica Kurok. var. antillarum
(Vain.) Kurok.
Floreana, Fernandina, Rdbida, Santa Cruz. Frequent on
trees, dead wood in the cactus forest, and occasionally on shore
rocks. Thallus ivory-textured, white with gray tones especially
on the essentially marginal isidia, white beneath. Never asso-
ciated with mosses but directly attached to the substrate.
Heterodermia comosa (Eschw.) Follm. & Redon, Willdenowia 6:446.
1972. Anaptychia comosa (Eschw.) Mass., A. podocarpa of re-
ports).
San Salvador: Taylor 916 (MO), Pike 56164; Santa Cruz:
40741, 41131, 44036. Never common, occurring scattered on small
twigs in the Scalesia forest area. Forming isolated rosettes
with ascending short laciniae and long white marginal cilia.
When sterile some lobes are cucullate and sorediate beneath; the
apothecia, when present, are pruinose and ciliate.
*Heterodermia corallophora (Tayl.) Skorepa, The Bryologist 75:
490. 1972. Anaptychia corallophora (Tayl.) Vainio.
San Cristébal, Lanier 63755;,Santa Cruz?.on.Psidium. 340706.
41145, Sipman 63493. Characterized by a soft foliose greenish
thallus with numerous laminal isidia. All material is sterile.
*Heterodermia lepidota Swinscow & Krog, Lichenologist 8:122-124.
plate 2.1976.
Isabela: Volcan Cerro Azul, S slope, above Iguana Cove,
250 m.s.m., on slanting trunk of Pisonia, Sipman 63504. Thal-
lus characterized by entire or dissected laminal squamules and/
or soredia or isidia (ours also has isidia). In contrast to
H. antillarum, the thallus is broader and forms large round
plaques instead of linear radiant lobes and it lacks the firm
ivory-white cortex.
*Heterodermia leucomela (L.) Poelt sens. lat. See Swinscow &
Krog, Lichenologist 8:134-129. 1976. (Physcia leucomela of
reports).
RAabida, Floreana, Isabela: Lich. Exs. COLO 508 (as Anap-
tychia stellata), 509 (f£. verrucifera), Pinta, San Salvador,
Santa Cruz. Locally abundant on trees and shrubs in the moist
zone. Long narrow ribbons, white above, white beneath, with
long black cilia; thallus straight or circinate at the apex;
sometimes isidiate. Until detailed microchemical study is made
of these collections it seems best to lump them under H. leuco-
mela. Numerous segregates have been proposed. Lich. Exs. 504,
distributed as Anaptychia circinalis (Zahlbr.) W. A. Weber, is
one such, having slender, terete branches and circinate branch
471
tips. This grows in sites with typical flat-laciniate H. leuco-
mela without intergradation.
*Heterodermia lutescens (Kurokawa) Follmann, Philippia 2:73.
17a
Isabela: Sipman 63549; Pinzén: Sipman 63581; Santa Cruz:
63487. The thallus characteristically has a lemon-yellow pig-
ment on the lower surface, and is sorediate near the lobe tips.
*Hypotrachyna costaricensis (Nyl.) Hale. Parmelia costaricensis
Nyl.
Santa Cruz: fern-sedge zone, summit area between Cerro
Crocker and El Puntudo, on Scalesia pedunculata, 63053. Thallus
gray, smooth and shiny, with long cylindric isidia; rhizinae
branched, underside black. No chemical reactions.
*Hypotrachyna microblasta (Vain.) Hale. Parmelia microblasta
Vain.
San Salvador: Pike 56194; Santa Cruz: El Puntudo, summit
grassland, 55396. Yellow, isidiate, rhizinae dichotomous, no
pseudocyphellae, maculae or fissures; med. white, Pd+ yellow, K+
yellowish, underside black.
*Lecanora caesiorubella Ach. ssp. glaucomodes (Nyl.) Imshaug &
Brodo, Nova Hedwigia 12:15.1966.
Santa Cruz: Common on Piscidia, cactus forest. Thallus
white; apothecia pruinose, slightly contorted; margin K+ yellow
to red (norstictic), Pd+ orange, (protocetraric), C-.
*Lecanora leprosa Fée. See Brodo (1984).
Floreana, RAabida, Santa Cruz, Wolf. A very common corti-
colous species on a variety of trees in the cactus forest; also
on mangroves. Superficially similar to Lecidea granifera but
with thinner apothecial margins, and with algae in the margin.
*Lecanora pseudopinguis W. A. Weber, Mycotaxon 13:102-103. 1981.
Endemic. Champion, Floreana, Isabela, Rabida, Plazas, San
Salvador, Santa Cruz: Lich. Exs. COLO 500 (Type). One of the
dominant lichens of the coastal rocks and seaward-facing bar-
rancos (one collection from 400 m.s.m.). Thallus and apothecia
dull yellow, C+ orange. Thalli at the type locality were so
badly damaged from fresh-water scouring during the 1982 El Nino
as to be unrecognizable.
*Lecanora sulfurescens Fée
Darwin, Plazas, San Salvador, Santa Cruz, Wolf. Common on
lava rocks near the coast but generally well back from the high
472
tide mark. Thallus crustose, white, with small, flat areolae;
apothecia from reddish brown to pale yellowish-brown, usually
immersed or nearly so, with a narrow rim.
*Lecidea chilena Zahlbr. L. aeruginosa Nyl. For description,
see Zahlbr. in Skottsberg, Acta Horti Goteb. 2:9. 1926.
Floreana: on lava, rocky point just south of Black Beach,
63013; Santa Cruz: on lava, Academy Bay, 40834. Disjunctive
from Chile. The green-pruinose apothecia are distinctive.
*Lecidea cf. granifera (Ach.) Vain.
Santa Cruz: 40082, 40197, 40337, 40423, 40475, 40616; San
Salvador: 40089, 41028. A white crust on various trees in the
cactus forest. The apothecia superficially resemble those of
Lecanora leprosa, but algae are absent from the thicker margin
and below the hypothecium. Our collections are not warty as
they usually are in Florida collections, and may represent a
segregate. It is highly unlikely that this taxon will remain in
Lecidea.
*Lecidea oreinoides (Koerb.) Weber & Hertel in Hertel, Herzogia
Pape Ve Shlt lee hi ab
Santa Cruz: 40039, 40067, 40860, 40862, 40867, 40871.
Frequent on boulders on shore and in the cactus forest. A ubi-
quitous lichen of the subtropics of both hemispheres. It has
received very many names, first under Aspicilia, which it su-
perficially resembles. Thallus white, cracky-areolate, with
black prothalline lines visible between the areoles; apothecia
round, totally immersed and occasionally running together.
*Leprocaulon tenellum (Tuck.) Nyl.
Fernandina, Floreana, Isabela, San Cristébal, San Salvador,
Santa Cruz: Lich. Exs. COLO 496 (as L. microscopicum). Common
on exposed crater rims often bathed in fog. Also present on the
mainland coast of Peru and Chile.
*Leptogium cyanescens (Ach.) Koerb.
Santa Cruz: Lich. Exs. COLO 506. On living fences in the
cultivated zone, and on the ground in summit heath. Thallus
slate gray, naked below, scarcely wrinkled, with cylindric to
flattened isidia on upper surface and margins.
*Leptogium isidiosellum (Riddle) Sierk, Bryologist 67:282-283.
1964.
Santa Cruz: on lava boulders, base of first barranco along
old trail to Bella Vista, lower edge of Transition zone, 40274.
Thallus brown to blue-gray, densely isidiate, with "pulled"
473
wrinkles beneath. The habitat, fide Sierk, is corticolous.
Leptogium marginellum (Sw.) S. Gray
Floreana, Isabela, San Salvador, Santa Cruz. Common on
trees in the moist zone. Characterized by marginal, light-
colored apothecia, surrounded by cylindrical clavate or lobulate
isidia; both sides of thallus distinctly sharp-wrinkled.
*Leptogium millegranum Sierk, The Bryologist 67:290. 1964.
Fernandina, San Salvador, Santa Cruz. Frequent on smooth-
barked trees in the cactus forest and lower Transition zone.
Thallus lead gray to greenish black, with alternating exceeding
ly thin, stretched areas and elongate high ridges covered with
small dark isidia.
*Leptogium punctulatum Nyl.
Santa Cruz: Scalesia pedunculata forest, north slope of
mMOUneweroOcker., G0 msm, 61462) Lich "Exs.('COLO' 505. “tPo M.
Joergensen. Forming densely tufted ascending or erect thalli on
horizontal branches of Scalesia on forest margin. Thallus
steel-gray, foveolate, broad-lobed, no tomentum or isidia; apo-
thecia common, sessile, amphithecium pale tan.
*Lobaria dissecta (Sw.) Ratisch.
Pinzon: mixed Scalesia forest of trees 4 meters tall, only
one thallus seen, on Scalesia branch, Sipman L-63580. Thallus
shining ivory-white above, tan, naked below; lobes with rounded
sinuses.
*Lopadium foliicola (Fée) R. Sant.
San Cristobal: on Citrus leaves below Tres Palos, Lanier
L-63763. !Vezda.
*Lopadium fuscum MUll.-Arg.
San Crist6ébal: on Persea americana leaves at Tres Palos,
Lanier 63700; on leaves of Coffea, Horneman farm, 40431. !Vezda.
*Lopadium puiggarii (MUll.-Arg.) A. Zahlbr.
Santa Cruz: 40216 p.p., 40264, 40265, 40266, 40435, 63331.
!Vezda. On leaves of various plants (Miconia robinsoniana,
Pteridium aquilinum) in the moist zone and summit heath.
*Melanotheca cruenta (Mont.) Mull.-Arg.
San Salvador: on twigs, Pike 54809. Thallus thin, smooth,
stained a dull red-orange, darker where the clusters of raised
perithecia occur.
474
*Myriotrema compunctum (Ach.) Hale, Mycotaxon 11:133. 1980.
Leptotrema compunctum (Ach.) MUll.-Arg. See Redinger (1936).
San Salvador: Pike: 56165:\\Santa* Cruz: 4034/7; 40357.) fich.
Exs. COLO 643, !Hale. Forming large gray unbroken sheetlike
crusts on Bursera in the cactus forest; apothecia immersed,
minute, the disk dilated, pruinose.
Myriotrema olivaceum Fée
San Salvador: on Xanthoxylum, 530 m.s.m., Pike 55414.
!Hale. Thallus smooth, conforming to the bark, pale tan,
apothecia in the depressions, in groups, not raised, the
openings minute (0.1 mm).
*Myriotrema wightii (T. Tayl.) Hale, Mycotaxon op. cit. p. 135.
Leptotrema wightii (Tayl.) MUll.-Arg.
Isabela: Cerro Azul, Sipman 63506; Santa Cruz: 40539. On
Pisonia and Bursera, often on dead wood. Thallus gray, thick,
continuous; apothecia immersed, punctiform, not raised, visible
as black-brown dots.
*Niebla sp. nov. ined. Ramalina ceruchoides H. Magn. ined.
Pinz6n: 375 m. on vertical face of big boulder, Sipman
63573. Extremely fragile. Thallus terete and slender, yel-
lowish, the branch tips with exposed medulla or incipient so-
redia. Although this is undescribed, it is important for the
record, since this material matches Santesson 1645, from Chile:
Prov. Atacama: Copiapo, on rocks in the desert NW of the town,
9.vii.1940. This is not the same plant as Niebla ceruchoides
Rundel & Bowler, ined., from California, which is not at all
fragile, esorediate, and green.
*Normandina pulchella (Borrer) Nyl.
Santa Cruz: (in packet with Byssoloma subdiscordans) 40250;
San Salvador: 56158. On leaves of Eugenia jambos and Xanthoxy-
lum, moist zone.
Ochrolechia pallescens (L.) Mass.
Floreana, Pinzén, San Cristébal, San Salvador, Santa Cruz,
Santa Fé. Warty gray crust with large thick-margined apothecia;
disk pinkish.
*Pannaria pannosa (Sw.) Del. Parmeliella pannosa (Sw.) MUull.-
Arg.
Isabela: Cerro Azul, south slope above Iguana Cove, 250
m.s.m., on a slanting Pisonia trunk, Sipman 63501. Thallus
sterile, densely isidiate except for the very margin, black
pannose-rhizinate below.
475
*Parmentaria astroidea Fée
Santa Cruz: on Ochroma (balsa) trees in plantation at Bella
Vista, 54837. Resembling Pyrenula nitida but perithecia grouped
and with a paler central wart, in section showing this to be a
focus of canals running from the various perithecia.
*Parmotrema conformatum (Vain.) Hale. Parmelia conformata Vain.
Floreana: on Bursera, trail from Black Beach to highlands,
62917. !Hale. Thallus yellow, black beneath, with incipient
soredia on ridges and margins.
*Parmotrema crinitum (Ach.) Choisy. Parmelia crinita Ach.
San Salvador: 56192; Santa Cruz: Academy Bay, on Bursera,
40622a, 40625a (with P. endosulphurea). Thallus ciliate, isi-
diate, black beneath margin brown, bare; med. K+ yellow, C-.
*Parmotrema cristiferum (Tayl.) Hale. Parmelia cristifera Tayl.
(Parmelia perlata and latissima of reports).
Fernandina, Floreana, Santa Cruz. Very common on smooth
barked trees, cactus forest through transition and moist zone.
Thallus with sorediate margins, no marginal cilia, cortex K+
yellow, medulla K+ red, Pd+ orange-red, C-, K-.
*Parmotrema dilatatum (Vain.) Hale. Parmelia dilatata Vain.
Santa Cruz: on rocks just above high tide mark, 40057,
40820, 40873 !Hale. Thallus gray-white, sorediate on upper
side at lobe tips, no cilia, cortex K+ yellow, medulla white,
K-, Pd+ yellow-orange, C-.
*Parmotrema dominicanum (Vain.) Hale. Parmelia dominicana Vain.
Mvoreana 629125 62913" “Santa Cruz, 40172-47186, "40890
Lich. Exs. COLO 136. !Hale. On Laguncularia and smooth-barked
trees of the cactus forest at low altitudes. Soralia greenish
(usnic acid), marginal on upturned lobes, cortex K+ yelow, Pd+
orange.
*Parmotrema eborinum (Hale) Hale. Parmelia eborina Hale.
Pinzén: caldera rim, 300 m.s.m., 40237, 40505. !Hale.
Thallus lacking soredia or isidia, ashy-white, often mono-
phyllous, fragile when dry, K+ yellowish, KC+ red, Pd+ orange.
*Parmotrema endosulphureum (Hillm.) Hale. Parmelia endosul-
phurea Hilln.
Pinzén: 40236, Stewart 3367 (FH, as P. latissima); San
Salvador: Stewart 1004 (FH, as P. latissima). Santa Cruz:
40120, 40239, 40282 p.p., 40624, 40625. On Bursera, Pisonia,
476
and other smooth-barked trees in the cactus forest. Thallus
eciliate, isidiate, medulla pale yellow.
*Parmotrema hypotropum (Nyl.) Hale. Parmelia hypotropa Nyl.
Santa Fé: Orr 40415. !Hale. Thallus ciliate, cortex
maculate, med. K+ red (atranorine and norstictic).
*Parmotrema leucosemothetum (Hue) Hale. Parmelia leucosemotheta
Hue
Pinzé6n: Cavagnaro 40151, 40159, 40494. On Croton bushes,
summit of the island. Thallus marginally sorediate, ciliate,
maculate, with pale blotches underside at margin, Med. K+ yel-
low-red (salacinic and atranorine), P+ pale orange-red, C-, KC-,
cortex K+ yellow.
Parmotrema peralbidum (Hale) Hale. Parmelia peralbida Hale
Floreana: 62905;..Santa Cruz: 40413, 40545, 63065. ‘On Bur-
sera in the cactus forest. A rather small Parmotrema, tightly
adnate; thallus white, soft and fragile, with many minute isid-
ia, becoming + sorediate; cortex K+ yellow, med. K-, C-, KCt+
rose (fleeting), Pd+ orange (atranorine and protocetraric).
*Parmotrema praesorediosum (Nyl.) Hale. Parmelia praesorediosa
Nyl.
Pinzé6n: 40499, 44017; Santa Cruz: 40446, 40542, 40621.
!Hale. On smooth-barked trees in the cactus forest, only rarely
on rock. Thallus white, eciliate, soralia marginal, forming
elongate, sinuous crescentic areas; noticeably thinner in
comparison with P. cristifera.
*Parmotrema reticulatum (Tayl.) Choisy. Parmelia reticulata
Tayl. |
Pinzén: 50517; San Salvador: Pike 55194; Santa Cruz: 40511,
40540, 55398. On trees (Xanthoxylum and Croton) and rocks in
the transition zone and summit grassland. Thallus with fine
reticulate cracks, ciliate, with laminal and marginal soredia;
med. K+ yellow-red (atranorine).
Parmotrema tinctorum (Nyl.) Hale. Parmelia tinctorum Nyl.
Isabela, Pinzén, San Salvador, Santa Cruz, Lich. Exs. COLO
114. Wolf. Abundant on smooth-barked trees in the cactus
forest, occasionally on Jasminocereus and rocks. A large Par-
motrema with pearl gray thallus with broad rounded lobes, simple
or branched laminal isidia; medulla C+ red.
|
|
*Parmotrema ultralucens (Krog) Hale. Parmelia subcrinita Nyl. ©
Floreana: 62915, Taylor 868 (MICH); Pinzén: 40491, Stewart —
'
477
BA9°p.p. (FH, as P. latissima); Santa Cruz: 40011,..40292, 63059.
!Hale. Wide ranging from cactus forest to summit heath, on
trees and rocks. Thallus not maculate, ciliate, isidiate, isi-
dia simple; lower side black with marginal naked area; medulla
white, C-, K+ red (salazinic).
*Parmotrema xanthinum (MUll.-Arg.) Hale. Parmelia xanthina
(Mi11.-Arg.) Vain.
Pinzé6n: caldera rim, 300 m.s.m., Cavagnaro 40757. Sub-
strate not indicated. Thallus yellow (usnic), cortex K+ yellow,
ciliate, med. white, underside naked at margin, medulla non-
reactive.
*Peltigera dolichorhiza (Nyl.) Nyl.
Isabela: Cerro Azul, south slope above Iguana Cove, 800
m.S.m., on steep mossy bank among ferns, v. d. Werff 64414,
64415.
*Peltigera erumpens (Tayl.) Elenk.
Isabela: Cerro Azul, 750 m.s.m., Sipman L-52; Santa Cruz:
on bare soil, summit heath, 40302, 41144. Gray-brown squamules
with erumpent patches of gray soredia in the center. Vitikai-
nen, in Poelt & Vezda (1981) treating P. spuria, evidently does
not include a esorediate form (P. erumpens [Tayl.] Elenk., P.
hazslinszkyi Gyelnik) in his species concept. P. didactyla
(With.) Laundon is the correct name for the fertile form of P.
spuria. I am not convinced that it is related to the sorediate
plant.
*Peltula euploca (Ach.) Wetmore
Champion: 62786; Darwin: Dawson 40900, 40901; Santa Cruz:
40071; Wolf: Cavagnaro 40917. A very inconspicuous lichen of
lava boulders near the shore. Thallus consisting of a single
thick squamule the color of the rock; margin gray, roughened.
Pertusaria albinea Tuck., Proc. Amer. Acad. 12:177. 1877.
Endemic. Type from "Galapagos Islands", Rev. T. Hill
(Hassler Exped.). Santa Cruz: 40346 appears to be a match for
the type. On Bursera. A smooth white species with low hemi-
spherical warts with a small dark ostiole; no soredia or isidia.
Pertusaria bispora (Farlow) Linder, Proc. Calif. Acad. Sci. 21:
213-214. 1934.
Genovesa, San Salvador, Santa Cruz. Thallus thin, smooth,
yellow, the ascocarps forming low mounds. Yellow Pertusaria
resembling this are common around the Pacific Basin from Cali-
fornia to Australia. It is unlikely that this name will be the
478
oldest one available. P. pustulata (Ach.) Duby seems to be a
likely possibility.
*Phaeographis dendritica (Ach.) MUll.-Arg.
Fernandina, Floreana, San Cristébal, Santa Cruz, San Sal-
vador. Common on Bursera and other smooth-barked trees in the
cactus forest. A distinctive species. Thallus white, exten-
sive; apothecia lirelliform, radiating, gray-pruinose, tapered
at the ends.
*Phyllopsora corallina (Eschw.) MUll.-Arg.
Floreana: on Psidium guajava, 62909; Santa Cruz: common on
living fences of Jatropha curcas, Horneman Farm, 40262. Thallus
of narrow green isidia standing on a mat of loosely interwoven
brown hyphae prominent around the thallus margin; apothecia
immature in ours, brown, biatorine.
*Physcia callosa Nyl.
Santa Cruz: 40013, 40343, 40198, 40449. Rare or infrequent
on lava blocks of barranco slope just north of Darwin Station,
vicinity of seismic station. Thallus white, finely dissected
and loosely attached, with coarse round soredia arising from the
underside of the lobe edges.
*Phaeophyscia hispidula (Ach.) Esslinger
Isabela: Cerro Azul, south slope above Iguana Cove, 250
m.s.m., on Pisonia, Sipman 63502, 63503. !Esslinger. Thallus
gray, with round greenish soralia and a dense mat of black stiff
rhizinae visible at the margins.
Physcia insularis A. Zahlbr., Ann. Mycol. 29:86. 1931.
Endemic. Floreana: Post Office Bay, Herre (Type in Vien
na); Santa Cruz: 30434, 40198, 40389, 40449. Zahlbruckner de-
scribed this species as being sorediate and the medulla K-. The |
cited specimens agree with this. Thomson (1963 p. 44) wrote:
"Although P. insularis is described as being sorediate, isotypes
of the original Galapagos Island material (Krypt. Vind. 3170)
have been examined in the U. S. National Herbarium and the her-
barium of the University of Colorado. These are not sorediate
but have the heavy coat of pruina of [P. biziana]." But the
exsiccati did not claim to be the type collection, only "ex loco
classico"! Thomson never saw the type, which Zahlbruckner did
not identify by number. Both P. insularis and an esorediate
species with pruinose apothecia and medulla K+ yellow (here
identified as P. mexicana B. de Lesd.) are abundant on Bursera
at Post Office Bay and the exsiccati may in fact consist of a
mixture with various packets having one or the other species.
479
*Physcia mexicana B. de Lesd.
Floreana: Post-office Bay, 54827; San Salvador: James Bay,
Dawson 41021. Thallus white, esorediate, the lobes narrow and
convex; apothecia pruinose; medulla K+ yellow.
*Physcia lacinulata MUll.-Arg.
Fernandina: west side of island, 320 m.s.m., on Bursera,
Cavagnaro 40231. !Esslinger. This is the first collection from
south of Costa Rica and the first in fruiting condition since
the type. Thallus tan-ash-brown, upper side K-, epruinose, with
numerous crowded-squamulose isidia or microlobules; marginal
lobes short, flabellate.
Physcia sorediosa (Vain.) Lynge
Fernandina: 40471; Floreana: Taylor 858 (specimen not found
at MO), reported by Dodge (1936); Santa Cruz: 40240-40242. A
large Physcia, white or yellowish, with closely contiguous plane
marginal lobes; soredia coarse, in globular laminal soralia,
becoming so numerous as to obscure the thallus surface;
apothecia usually present, with prominent smooth rim; cortex and
medulla K+ yellow; underside black.
*Physma byrsinum (Ach.) MUull.-Arg.
Isabela: Cerro Azul, on Scalesia trunk, south slope above
Iguana Cove, 350 m.s.m., Sipman 63535. Thallus thick, swelling
when wet, ashy-gray, much wrinkled and foveolate when dry; apo-
thecia numerous, large, with reddish-brown disk and wrinkled
gray margin.
*Pseudocyphellaria argyracea (Del.) Vain.
Pinzén: 450 m.s.m., on shrubs on exposed rock ridge with
ferns and scattered shrubs, Sipman 63582. Thallus brown with
white dorsal pseudocyphellae developing clusters of brown isid-
ia; underside brown-tomentose with large white pseudocyphellae.
Pseudocyphellaria aurata (Ach.) Vain.
Fernandina, Floreana, Isabela, Pinzén, San Cristébal,
Santa Cruz, San Salvador. Infrequent on trees and shrubs in the
moist zone, not as common as its collections indicate, since it
is taken wherever encountered, because of its striking appear-
ance. Thallus gray with yellow underside and yellow pseudo-
cyphellae; soredia marginal, yellow. Some specimens must differ
chemically, for on drying some soon turn dull brick red while
others remain gray. This does not seem to depend in the manner
of desiccation.
480
*Pseudocyphellaria crocata (L.) Vain.
San Salvador: Pike 56162; Santa Cruz: Table Mt., 40514,
41184. Less frequent than the last. Thallus green-brown,
underside tan, tomentose, with yellow pseudocyphellae; upper
surface with marginal and laminal yellow soredia.
*Pseudocyphellaria mougeotiana Vain. var. aurigera (Del.) Vain.
P. xantholoma (Del.) Dodge.
Floreana: Itow 40719, 40758; San Salvador: Pike 56182;
Santa Cruz: Table Mountain, 40487. Infrequent, on trees and
shrubs in the moist zone. Thallus shiny brown with elongate
marginal and laminal gray soralia, eroding to expose a yellow
medulla, dark tomentose beneath, naked and pale brown near the
margins.
*Pseudoparmelia caroliniana (Nyl.) Hale. Parmelia caroliniana
Nyl.
Isabela: Sipman 63538, 63547; Santa Cruz: 40384, 44028,
44031; San Salvador: Pike 55210, 56488. On various trees in the
moist zone. Thallus yellowish-green, isidiate or pustulate, the
isidia narrow and elongate, tending to be in clusters and often
branched; cortex surface minutely ridged or wrinkled.
*Pseudoparmelia leucoxantha (MUll.-Arg.) Hale
Floreana: trail from Black Beach to highlands, 62892.
'Hale. On rocks; thallus yellow, laminally and marginally
sorediate, black beneath; sterile.
*Psoroglaena cubensis MUll.-Arg., Flora 74:381. 1891.
Reported by Dodge (1936), from Hood: on rock, islet,
Gardner Bay, Taylor 872. The specimen (FH) is a species of Pel-
tula. However, I have collected the real Psoroglaena cubensis
and report it here. Santa Cruz: on old fern rhizomes, base of
the first barranco behind the Darwin Station, in shaded talus,
40298. Sterile. A minutely foliose olive green thallus, irreg-
ularly dichotomous-pinnately divided. The type collection was —
also collected on fern rhizomes, possibly an obligate substrate.
COLO 146, San Salvador. Common on smooth barked trees and
branches, aerial roots of mangroves, etc., mangrove zone,
coastal scrub and lower cactus forest. A bright orange thin
crust with minute perithecia in surface depressions.
i
|
Pyrenula cerina Eschw. P. aurantiaca Fée |
Genovesa, Isabela, Marchena, Rdbida, Santa Cruz: Lich. Exs.
*Pyrgillus javanicus Nyl.
481
Santa Cruz: on decorticate, termite-worked wood of dead
leaning Bursera tree, cactus forest just north of Darwin Sta-
tion, 30 m.s.m.
Pyxine berteriana (Fée) Imshaug
Santa Cruz: 40355, 63281. On Bursera in the cactus forest.
Thallus with radiating plane or concave lobes, smooth, not pru-
inose, flabellate-lobed, esorediate, UV+ yellow; medulla pale
yellowish; apothecia black, sessile, hypothecial stipe absent.
Compares well with Kalb: Lich. Neotropici 194.
*Pyxine caesiopruinosa (Tuck. in Nyl.) Imshaug
Santa Cruz: on talus of first barranco near seismic station
behind Darwin Station, 40876. Thallus dark gray, paler near the
margins, with small capitate laminal soralia; medulla where
broken showing through as pale salmon color; UV+ yellow.
Compares exactly with Imshaug 24790, from Cuba.
*Pyxine cocoes (Sw.) Nyl.
Plazas: 40600; Santa Fé: 40428; San Salvador: Pike 55407.
Thallus white, often with thick pruina; soredia copious on lam-
ina and margin; thallus not reticulate, medulla white; hypo-
thecial stipe K+ red; UV+ yellow.
*Pyxine connectens (Vain.) Malme
Floreana: 62907 (matched with the holotype), Isabela,
Pinta, San Salvador, Santa Cruz, Wolf. Common on Bursera,
Acacia, and occasionally Opuntia. Thallus thick, plane,
marginally so rediate, often the older parts cracked, the
younger lobes with a crustlike pruina, cortex K-, UV+ yellow.
*Pyxine eschweileri (Tuck.) Vain.
Floreana, Pinzén, Santa Cruz. Equally common on trees and
rocks in the coastal and cactus forest zone. Similar in ap-
pearance to P. connectens but thallus UV-; medulla somewhat
yellowish.
*Pyxine pringlei Imshaug, Trans. Amer. Microscopic Soc. 76:263.
O57)
Floreana, Santa Cruz: Lich. Exs. COLO 111; Wolf. Common on
lava rocks in the cactus forest. Thallus pale yellow with black
plane apothecia; stipe reddish, K+ purple. UV+ yellow.
*Ramalina anceps Nyl.
Rabida: Stewart 421 (as Alectoria sarmentosa). !Rundel,
2975. Similar to R. usnea but with salazinic and norstictic
acids (R. usnea lacks norstictic acid).
482
Ramalina complanata (Sw.) Ach.
Santa Fé: Orr 40416; Wolf: Schuster 40230. !Rundel. On
coastal shrubs. Thallus rigid, flattened, foveate and longi-
tudinally ridged, containing usnic and divaricatic acids.
*Ramalina denticulata (Eschw.) Nyl.
Espaniola, Genovesa, Isabela, Plazas, San Cristébal, Santa
Cruz, Lich. Exs. COLO 147 (as R. complanata), Santa Fé, San
Salvador, Tortuga, Wolf. All !Rundel. Similar to R. complanata
and usually so reported, but with salazinic acid.
*Ramalina furcellata (Mont.) Zahlbr.
Floreana, Pinzén, Radbida, San Salvador, Santa Cruz. On
coastal shrubs. Thallus with basal parts flattened, branches
terete, erect or somewhat pendulous, with soredia arising from
pseudocyphellae; medulla Pd+ yellow; very fragile when dry.
*Ramalina linearis (Sw.) Ach.
Floreana, Isabela, Santa Cruz, Santa Fé. Thallus narrow,
thin, deeply and often connivent-canaliculate, with marginal
apothecia; soredia and isidia lacking; med. K+red, Pd+ yellow.
*Ramalina peruviana Ach.
Floreana: 40777; Santa Cruz: 40286, 44035, 40697; San Sal-
vador: Pike 55419. All !Rundel. On coastal rocks, and also on
shrubs in the transition and moist zones. Thallus almost terete
but somewhat angular and striate, cracked on the edges and with
small soralia; sekikaic and ramalinolic acids. Several speci-
mens, thought to be this species, were held aside by Rundel be
cause they contain salazinic acid.
*Ramalina sorediantha Nyl. |
Santa Cruz: Lich. Exs. COLO 137. Common on Jatropha curcas—
living fences, Bella Vista. Thalli forming small balls, lacin-
iae slender, canaliculate, with ball-like soralia on the branch ©
tips; medulla K+ red, Pd+ yellow. i
Ramalina usnea (L.) Howe |
Espafiola, Floreana, Pinzén, Rabida, San Cristébal, Santa :
Cruz: Lich. Exs. COLO 115, Santa Fé, San Salvador... An extremely
abundant species in the transition and moist zones, festooning |
trees wherever fog persists. Probably some of the material will
prove to belong to R. anceps; the lines separating these taxa
are not clear to me.”
Roccella babingtonii Mont. R. intricata, R. peruensis, of pre-
483
vious reports.
Baltra, Bartolomé, Daphne Major, Espafiola, Floreana, Geno-
vesa. lsabela, Pinta, Pinz6én, Rabida, San) Cristé6bal, Santa Cruz:
Lich. Exs. COLO 109, Santa Fé, Wolf. Extremely abundant on
often moribund trees (Bursera, Acacia) and coastal shrubs. One
of the commercial sources of orchil, formerly collected on the
islands for commercial exploitation of the dyestuff by factories
on the mainland. Thallus gray, flattened, with broadly linear
branches, sorediate; cortex C+ red (lecanoric acid).
*Roccella galapagoensis Follmann, Nova Hedwigia 15:337-340.
1968. R. portentosa, incorrectly, of reports.
Baltra, Bartolomé, Darwin, Espafiola, Floreana, Gardner,
Isabela, Pinzén, Santa Fé, San Salvador, Santa Cruz: Lich. Exs.
COLO 112, Type; Wolf. Locally abundant on coastal rocks.
Thallus terete, to 5 mm thick, forming long ropy strands
ascending at the ends; apothecia immersed, round, black.
Chemically vari able, containing erythrin, galapagin, portentol,
protocetraric acid, roccellic acid.
*Roccellina badia Tehlér, Opera Bot. 70:67. 1983.
Floreana: 62896; San Salvador: 56478: Santa Cruz: 40824,
62854. On massive lava blocks usually vertically exposed.
Thallus thin, minutely areolate, creamy white with a purplish
cast, occasional point soredia scattered; apothecia tan, with
concolorous rim. Cortex C+ red. Known only from Galapagos and
adjacent coastal Peru.
Schistophoron tenue Stirton
santa Cruz: Herre 41158, Lich. Exs. COLO 495: Isabela,
Sipman 63512. On smooth-barked trees (Pisonia), once collected
(Isabela) on decaying plant bases. Collected at Academy Bay in
1929 by Herre, I did not rediscover this species until 1976,
when I found an entire tree trunk covered with it. For many
years known only from Galdpagos and west Africa, it recently has
been discovered on the Ecuadorean mainland. Thallus smooth,
white; fruiting structures dome-shaped, sometimes curved and
slightly elongate, with a black, simple or sometimes forked
median line. |
*Sphinctrina podocarpa Mull. -Arg.
San Salvador: on Pertusaria, in Psidium-Bursera forest on
pahoehoehoe lava flow near salt lake on James Bay, 170 m.s.m.,
Pike 56178. Podetia nine-pin-shaped, red-brown with bluish
base, 0.5 mm high.
*Spilonema revertens Nyl.
484
Santa Cruz: on lava blocks along old trail from Puerta
Ayora to Bella Vista, 63304. Forming inconspicuous black balls
a few mm diam, in and around crevices of the rock. Thallus
minutely fruticulose, the base distinctly bluish where separated
from the substrate.
*Sporopodium leprieurii Mont.
Santa Cruz: on leaves of Miconia robinsoniana, summit
area, 40213. !Santesson.
*Stereocaulon azulense Yoshimura & W. A. Weber, sp. nov. See
Appendix III.
*Stereocaulon microcarpum MUll.-Arg.
San Cristobal: El Junco, Lanier 63748, 63761. On lava
rocks.
*Stereocaulon weberi Lamb, J. Hattori Bot. Lab. 43:258. 1977.
Isabela: 75985, Sipman 63528; San Cristébal: El Junco
(mixed with S. microcarpum), Lanier 63748 p.p., 63762 p.p.; |
Santa Cruz: Lich. Exs. COLO 494 (Type, distributed as S.
microcarpum), 63061, 63038. On lava, summit areas of the higher |
islands. Thallus erect, little-branched, with granulose-soredi- |
iform phyllocladia. |
*Sticta filix (Sw.) Nyl.
Santa Cruz: 40509, 40512, 44027, 55388, 55402; San Salva- |
dor: Pike 55422. On Scalesia trees in the moist zone. Thallus
monophyllous, fern-like, erect, green when wet, brown beneath
with prominent "veins", finely laciniate with a thickened basal |
holdfast, the rhizinae very short.
*Sticta fuliginosa (Dicks.) Ach.
San Salvador: in dense stand of Xanthoxylum and Psychotria
between James Bay and summit, 780 m.s.m., Pike 55421. Thallus |
monophyllous, gray with broad rounded lobes, above with very
fine black cylindrical isidia in clumps; underside pale. :
*Sticta sylvatica (Huds.) Ach.
Santa Cruz: 40430; San Salvador, Pike 55423. On Miconia
and other shrubs. Thallus gray, with rounded lobes, surface
with scattered lead-gray isidia, underside pale, with pale
rhizinae. Our specimens very small and fragmentary.
Sticta weigelii (Isert ex Ach.) Vain.
Isabela, San Salvador, Santa Cruz. Omnipresent species on
trees and shrubs in the moist zone. Thallus brown to purplish-
RR arn —— >
485
black. margins (and often lamina) isidiate, smooth (or in forma
peruviana [Del.] Vain.) depressed-reticulate.
*Strigula nitidula Mont.
Santa Cruz: on leaves of Eugenia jambos, along trail to
Horneman place just above Bella Vista, 64131 p.p. (with Bysso-
loma subdiscordans). !Vezda.
*Strigula subtilissima (Fée) MUll.-Arg.
Santa Cruz: on leaves of Eugenia jambos, along trail to
Horneman place just above Bella Vista, 64131 p.p. (with Bysso-
loma subdiscordans). !Vezda.
*Tapellaria epiphylla (MUll.-Arg.) R. Sant.
San Cristébal: on Citrus leaves below Tres Palos, Lanier
63764. !Vezda; Santa Cruz: on leaves of Eugenia jambos, along
trail to Horneman place above Bella Vista, 42047 (with Byssoloma
subdiscordans). !Vezda.
*Tapellaria nana (Fée) R. Sant.
Floreana: 64132; San Cristébal: Gradstein 63825; Santa
Cruz: 40211, 40431. On Eugenia, Coffea, Citrus and Miconia,
moist zone. !Vezda.
*Tapellaria nigrata (MUll.-Arg.) R. Sant.
San Cristébal: 63763 p.p. (with Lopadium foliicola), 63769.
!Vezda. On Citrus and mango leaves.
Teloschistes flavicans (Sw.) Norm.
Espanola, Floreana, Isabela, Pinzén, Rabida, San Cristobal,
San Salvador, Santa Cruz: Lich. Exs. COLO 108; Santa Fé, Wolf.
Common on twigs in the cactus forest, infrequent in moist zone.
Thallus fruticose, orange, terete, with slit-soralia, usually
sterile; apothecia orange with thalline rim.
*Tephromela atra (Huds.) J. Hafelln. Lecanora atra (Huds.)
Ach.
San Salvador: on Bursera, Pike 56155 (cortex UV-, medulla
UV+). The genus is presently under study by Hafellner.
*Trapelia coarctata (Sm.) Choisy. Lecidea coarctata (Sm.) Nyl.
Santa Cruz: base of El Puntudo, west of Cerro Crocker,
55385. The only common and well-developed crustose lichen on
the almost constantly wet rocks of the summit grassland.
Thallus of minute scattered white squamules.
*Trapeliopsis granulosa (Hoffm.) H. T. Lumbsch. Lecidea granu-
486
losa (Hoffm.) Ach.
Santa Cruz: on plant debris, saddle between El Puntudo amd
Cerro Crocker, 700 m.s.m., 63337.
*Tricharia vulgaris (MUll.-Arg.) R. Sant.
Santa Cruz: on leaves of Coffea, Horneman garden, 40433
p-p. (with Aspidophyllum fugiens). !Vezda
*Trypethelium eluteriae Spreng.
Santa Cruz: 40410; San Salvador: Pike 56161. Common on
Bursera in the cactus forest. Thallus pale, gray or yellowish;
stroma high, rounded, more or less constricted at base, dis-
crete, with numerous punctiform ostioles; interperithecial areas
yellow, K+ violet.
*Trypethelium mastoideum Ach.
Fernandina: Cavagnaro 40476, 40484; San Salvador: Pike
56189; Santa Cruz: 40409, 54833. Common on smooth-barked trees
(Bursera, Ochroma, Xanthoxylum). Thallus smooth, yellowish, the
stroma dark, strongly contrasting, raised but not constricted at
base, with numerous punctiform ostioles; perithecia black, but
yellow areas tending to turn orange.
*Trypethelium ochroleucum (Eschw. in Mart.) Nyl.
Fernandina: Cavagnaro 40478; Santa Cruz: 40051, 40411,
40557, 40612, 54824; San Salvador: Pike 55226. Common on Bur-
sera, Psidium, Piscidia, Xanthoxylum in the cactus forest.
Thallus yellowish, covering large areas of bark as if painted
on; perithecia in clusters in slightly raised stromae, not
constricted at base, the ostioles black.
*Trypethelium tropicum (Ach.) Mull.-Arg. Zignoella magnoliae
Tracy & Earle, Z. lichenoides von Hoehnel, Z. nobilis Rehn,
acc. to Rogerson (1965 in litt.). Possibly not a lichenized
fungus.
Floreana, 62990; Santa Cruz: on dead termite-ridden tree,
moist zone below Bella Vista, 40209, 40341, 41200. On Bursera,
Citrus, Jatropha curcas fences, moist zone. Thallus smooth,
pale; fruits cannon-ball-round, black, in chains, with prominent |
pale ostiole. |
*Usnea antillarum (Vain.) Zahlbr.
Fernandina: Cavagnaro 40441; Pinzén: Cavagnaro 40050,
40518; Santa Cruz: 63311, Stewart 404; Santa Fé: Koford 40790.
Stout, branched from the base, green to pale green, cortex and
medulla very thin, the axis very thick, excavate inside; soredia |
arising commonly along the smaller branches as thin eruptive
487
sites, the soredia soon becoming isidiate and spinuliform, some
times the thallus wholly isidiate.
Usnea arthroclada Fée
Fernandina: Snodgrass & Heller (FH); Isabela: Stewart 408;
Pinta: D. Weber 55223; San Salvador: Pike 56190. Infrequent on
Bursera in the Transition zone. Thallus long, pendulous,
terete, annularly cracked, lacking papillae, soredia, or isidia;
stems sparsely dichotomous, the branches closely parallel so
that the stems appear simple; no pseudocyphellae; main branches
with many perpendicular short branchlets; medulla K+ red.
*Usnea cladocarpa Fée
Espanola: Itow 40735, 40769; Floreana: Itow 44003; Santa
Cruz 9200/73, Lich. Exs. COLO 507. On coastal shrubs and Bur-
sera. Thallus less than 3 cm, little branched, stout, with
thick stems (1.5-2.0 mm), papillose and ciliate; apothecia cup-
like, ours with short cilia; soredia absent; plants sometimes
degenerate and foveate, medulla K-.
*Usnea paradoxa (Zahlbr.) Mot.
Santa Cruz: Kastdalen 40029; Lich. Exs. COLO 366. On
shrubs in moist areas (the tortoise ponds near Las Casitas).
Thallus very long and essentially unbranched except for the
short secondaries; main stem sharply angular, 2 mm wide or more,
secondary branchlets sorediate; medulla K+ yellow becoming
blood-red.
*Usnea rubiginea (Michx.) Mass. U. rubescens Stirt.
Espanola, Floreana, Pinzén, Rabida, San Cristdébal, San
Salvador, Santa Cruz, Santa Fé. Thallus bushy-branched, cortex
with red spots or totally red; medulla white, K+ yellow becoming
red; branchlets with fine cylindric isidia and soredia. Fertile
specimens, not mentioned by Motyka, were collected on Pinzén:
40152, 40496, 41143, and Floreana: 40452, and are essentially
esorediate.
*Xanthoparmelia subramigera (Gyelnik) Hale. Parmelia subrami-
gera Gyelnik
Pinzén: Dawson 40904; Santa Cruz: 40858, 40923. !Hale.
Frequent on rocks in the cactus forest. Thallus yellow-green,
isidiate.
*Xanthoria candelaria (L.) Kickx
Isabela: 63261; Santa Cruz: very rare, on Piscidia,
40564; on Bursera, 40361. Thallus orange, forming appressed
sterile laciniae with sorediate apices.
488
REJECTED REPORTS, SYNONYMS AND MISAPPLIED NAMES
Alectoria sarmentosa Ach. A misidentification of Ramalina
‘usnea. See discussion in Weber (1966, p. 196).
Anaptychia leucomelaena: see Heterodermia leucomela
Anaptychia podocarpa (Bél.) Trev. Reported by Dodge
(1936). The specimens belong to Heterodermia comosa.
Arthonia cinnabarina (DC.) Wallr. Farlow (1902) = A. gre-
garia.
Arthonia nivea Willey, Synops. Gen. Arthonia, p. 5. 1890.
The holotype collection is "Galapagos, Hassler Exped."
(!US), on smooth bark, evidently Bursera. Since the "Hassler"
expedition covered a greater area than Galapagos and several
collections have been of species that for reason of non-volcanic
substrate could not possibly have come from the islands, it
seems best to delete this species from the list until verified
collections are forthcoming. Arthonia is a very common genus on
the islands, and I have many collections, but monographic study
is necessary for determinations.
Buellia umbrina Malme: Dodge (1936). The specimen is
Taylor 47874g (FH) in the packet of "Caloplaca muelleri". This
is a Buellia, and is matched by COLO 40860, Buellia indet.
Thallus brown, epithecium olivaceous, hymenium 40-50mu, IKI+
blue, K=, hypoth. pale brown, IKI+ blue, exciple brown to pale
within, spores brown, 2-celled, 9/x5mu, thick-walled with wide
septum; thallus Pd+ yellow-orange, medulla K+ yellow to red, C-,
Pd-, IKI-. Neither of these specimens matches Malme’s
description. The medulla in B. umbrina is K-.
Buellia xanthinula (MU11.-Arg.) Malme was reported by
Dodge (1936) from a specimen: Taylor 874d from Gardner Islet.
The specimen is Buellia straminea.
Caloplaca elegans (Link) Th. Fr. This was reported by
Dodge (1936) from San Salvador: Taylor 859 (MO in 1965). The
specimen is isidiate and belongs to Caloplaca isidiosa (Vain. )
Zahlbr.
Caloplaca muelleri (Vain.) Zahlbr. Reported by Dodge
(1936). The specimen (Taylor 874b) comprises the following:
Caloplaca isidiosa, Buellia straminea, Buellia "umbrina", Lecid-
ea oreinoides (not reported by Dodge although it dominates the
489
packet), Diploschistes actinostomus (reported by Dodge as Lepto-
trema species, immature), and a species of Dirinaria.
Caloplaca murorum (Hoffm.) Th. Fr. Reported by Stewart
(1912) as Placodium murorum, the specimen was undoubtedly C.
isidiosa, although it was not located at FH.
Caloplaca rugulosa (Nyl.) Zahlbr. Reported by Dodge (1936)
from Floreana, Taylor 859. The specimen could not be found
either at FH or MO. It should be rejected because of the gener
al unreliability of Dodge’s determinations.
Cladonia chlorophaea (Flk.) Spreng. Reported by Weber
(Lich. Exs. COLO 139). The specimens belong to C. balfourii.
Cladonia fimbriata (L.) Fr. Reported by Stewart (1912).
The specimens belong to C. pityrea.
Cladonia furcata f. adspersa Flk. Reported by Stewart
(1912). The specimens belong to C. pityrea.
Cladonia galapagosensis Ahti= Cladina confusa
Cladonia polia R. Sant. Reported by Weber (Lich. Exs. COLO
106). = Cladina confusa.
Cladonia pycnoclada, reported by Farlow (1902) and Stewart
(1912). The specimens are Cladina confusa.
Coenogonium sp. Reported by Stewart (1912). The specimen
is not Coenogonium but a green algal scum occurring on shore
shrubs.
Cora pavonia (Sw.) Fr. = Dictyonema montanum
Dictyonema sericeum. Reported by Howell (1942). The
specimens are D. guadeloupense.
Chiodecton sanguineum (Sw.) Vain. = Herpothallon sanguineum
Dirina herrei A. Zahlbr. = D. approximata.
Lecanora glaucovirens Tuck., Proc. Amer. Acad. 12:172.
1877. The Type: Hassler Expedition (FH) consists of four pieces
of smooth bark, each with a plaque. The thallus is greenish-
gray, grading at the margin to a distinctly pale prothallus; the
490
apothecia are pruinose. Microscopic analysis shows, however,
that this is not a Lecanora; the rim, is falsely lecanorine as
in Lecidea granifera Ach., but in contrast to the latter, the
hypothecium is hyaline. The spores are 10-12 x 6-/mu. Dispo-
sition of this taxon await study of the South American Lecideae.
In the absence of any later collections, I doubt that this
lichen really came from Galapagos.
Leptotrema mastoideum Mull.-Arg. The specimen (Taylor
903) from Floreana (!MO), reported by Dodge (1936) is Diplo-
schistes scruposus.
Pannaria molybdaea (Pers.) Tuck. = Coccocarpia palmicola.
Parmelia camtschadalis Eschw. The specimens reported by
Stewart (1912) belong to Everniastrum vexans.
Parmelia latissima Fée. Reported by Stewart (1912). The
specimens (FH) belong to Parmotrema tinctorum, P. subcrinita,
P. endosulphurea, and P. cristifera!
Parmelia perlata Kremp. Reported by Farlow (1902) and
Stewart (1912). The specimens belong to Parmotrema cristifera.
Parmelia soyauxii MUll.-Arg. Reported by Dodge (1936).
Th specimen was not found at FH or MO. This Dodge determination
is not to be accepted.
Pertusaria colobina Tuck. The type is a collection from the
Hassler Expedition. Santesson examined it and found that on the
four pieces of bark, marked a,b,c,d, were (a and d) Coccotrema
granulata (Hook. f. & Tayl.) R. Sant., and (b) Lepolichen cocco-
phorus (Mont.) Trev. It is doubtful, since a and b are labelled
"Galapagos" that there is any Pertusaria here at all, further
doubtful that any of the pieces came from Galdpagos. In the
packet is an annotation "Lepolichen colobinus (Tuck.) G. T.
Johnson". Tuckerman’s handwritten notes read: "sp 8.nae in
ellipt , simplice (Pertusaria) 46-56/23-25mu iis lich. u. i mm
similis. Hassler Exped. T. Hill". Santesson agrees that the
species present are distinctive Chilean species and could not
have been taken on Galdpagos. All Hassler Expedition "Gala-
pagos" collections, unless re-collected on Galapagos, should be
regarded as extremely doubtful records.
Physcia adglutinata (Flk.) Nyl. Reported by Dodge (1936)
from Isabela (Taylor 853). The specimen is a fragment of
Dirinaria sp., on coastal rock with bits of Buellia straminea].
a ee Se ee eee
491
Physcia aegialita (Ach.) Nyl. Reported by Dodge (1936). =
Dirinaria aegialita.
Physcia alba (Fée) MUll.-Arg. Reported by Dodge (1936)
from Hood (Taylor 8/4e). The fragment is in a packet of "Calo-
placa muelleri", and is a Dirinaria sp.].
Physcia aspera H. Magn. A variety of taxa have been re-
ported under this name. Stewart 357 (FH) was cited by Thomson
(1963), but it matches COLO 40110, Dirinaria caesiopicta.
Physcia leucomela (L.) Michx. = Heterodermia leucomela.
Physcia picta (Sw.) Nyl. Reports (Stewart 1912, Thomson
1963) are misdeterminations of Dirinaria applanata and D.
leopoldii.
Placopsis cribellans (Nyl.) Rasanen. Reported from a
Hassler Expedition collection. The specimens must have come
from Tierra del Fuego. See discussion in Weber (1966, p. 196).
Pseudocyphellaria xantholoma (Del.) Dodge = P. mougeotiana
var. aurigera.
Pyxine glebosa Tucks ~*Synops. N. “Amer. “Lich. 1279-71882.
"Galapagos Islands, Dr. Hill in Hassler Exp." Contrary to Ims-
haug’s annotation and his notes in Bryologist 58:283. 195, the
type specimen (!FH) is Pyxine and not Buellia. Superficially it
is similar to P. pringlei but UV-, there is a well-developed
pulpa or hypothecium as well as a brown stipe. The epithecium
and stipe in P. pringlei are K+ violet, in P. glebosa K-. The
spores are similar to those of P. pringlei (12-17[20]x5-7mu) and
mischoblastiomorphic. The apothecia are prominently enveloped
by a pseudothalloid margin. According to Imhaug’s treatment
this would come to P. endoleuca (MUll.-Arg.) Vain. However,
the questionable collecting data and the fragmentary condition
of the specimens argue against synonymizing a well-established
taxon. Nothing like it has been collected recently.
Ramalina dasypoga Tuck. The report is based on Taylor 911
(Dodge 1936). Taylor's collection (MO) comprises two packets of
R. furcellata and one of R. usnea.
Ramalina farinacea Ach. Reported by Dodge (1936). The
specimens represent a species, probably R. furcellata), with
terete thalli and globular soralia. Magnusson saw these
492
specimens but did not annotate them. Stewart 371 (FH) was
annotated by Magnusson as "R. complanata var."|]
Ramalina indica Fr. Reported by Farlow (1902) and Stewart
(1912) and Dodge (1936) from a specimen from Floreana (!FH)
collected by N. J. Andersson. The specimens fit my concept of
R. complanata sens. lat.
Ramalina interponens Nyl. The report by Dodge (1936) is
incorrect. The specimen, Taylor 873 (!MO), from Gardner Bay, is
R. complanata sens. lat. Chemical tests have not been made.
Ramalina puiggarii MUll.-Arg. All specimens reported by
Dodge (1936) belong to what I am calling R. linearis.
Ramalina subasperata Nyl. Reported by Dodge (1936). The
specimen represents R. complanata sens. lat.
Ramalina subfraxinea Nyl. Reported by Dodge (1936). The
specimen represents R. complanata sens. lat.
Ramalina usneoides (L.) Howe = R. usnea.
Roccella difficilis Darb. This taxon was suspected by
Darbishire to be a saxicolous form of R. babingtonii. The sub-
sequent report by Dodge (1936) is incorrect; the cited Taylor
specimen 854 (!FH, MO) is Roccella galapagoensis.
Roccellodea nigerrima Darb., Ann. Crypt. Exot. 5:153-158.
19324
The type of the genus and species came from "Galapagos",
J. Hill, Hassler expedition (herb. Darb., isotype FH). The
lichen is extremely similar to R. galapagoensis but was said to
have had the medulla of older parts black, and apothecia which
are irregular in shape (5x1.5 mm), and a colorless "hypothecium"
(dark in Roccella). Soredia were present. Two Stewart speci-
mens cited by Darbishire (1935) from Espafiola and Floreana are
sterile; how these could have been so determined is not clear.
Until more material is discovered the identity (and type locali-
ty) of this taxon remain a mystery. Perhaps somewhere in the
rich (and chemically variable) material of R. galapagoensis,
some of which is sorediate, may come an answer.
Sticta quercizans Del. Reported by Stewart (1912) = S.
weigelii.
Teloschistes exilis (Michx.) Vain. Reported by Dodge
LE
493
(1936). The specimen (Taylor 865, FH) is typical T. flavicans,
sorediate and sterile.
Usnea ceratina Ach. Reported by Farlow (1902) and Stewart
(1912). A common waste-basket assignment for unidentifiable
tropical Usnea. At least one collection, Stewart 405, is U. ru-
biginea. Herre noted on the specimens: "None of these remotely
resemble U. ceratina". A Baur specimen of U. rubiginea (FH) was
called U. barbata. Old identifications of Usnea, and many recent
ones, including my own, need to be taken with a grain of salt.
Usnea longissima Ach. Reported by Stewart (1912). The
specimens belong to U. amabilis.
Usnea plicata Hoffm. Reported on the basis of scrappy
specimens collected by J. H. Andersson and Charles Darwin.
Herre’s remarks could apply here as well.
APPENDIX II: NEW COMBINATION
Buellia mamillana (Tuck.) W. A. Weber, comb. nov. Rinodina
mamillana Tuck., Proc. Amer. Acad. 7:226. 1866 (publ. 1868).
Lecidea glaziouana Kremp., Flora 59:317. 1876. Buellia glaziou-
SpdeMullneAte (er woOra 69.19. 1880,
The type (!FH) was collected on Oahu, Hawaii, by Mann. The
Galapagos report by Tuckerman (1887) was based on a specimen
collected by Hill on the Hassler Expedition. The holotype (!FH)
in all respects agrees with Buellia glaziouana. Tuckerman
probably included this in Rinodina due to the false thalloid rim
of the apothecium. In section it is not lecanorine. The
epithet, mamillana, was chosen to denote a rugose-plicate
(teratological?) development of the apothecial disk. Tuckerman
himself attached little significance to this feature in pointing
out that the Galapagos material lacked it. Buellia glaziouana
has been described as a Rinodina several times, viz: R. thomae
Tuck., R. contiguella Vain., and also as Melanaspicilia.
Chemically, spot tests of both collections cited by Tucker
man give the thallus K+ yellow wash, Pd+ sordid yellowish, C-.
medulla IKI+ blue. Imshaug (1951) pointed out that the type of
B. glaziouana has an IKI+ blue reaction in the medulla, contrary
to statements of Vainio and Malme. Magnusson, in Cat. Lich.
Hawaii (1955 p. 386) points out that Rinodina mamillana Tuck.
has an IKI+ blue reaction in the medulla, contrary to Zahlbruck-
ner’s finding.
The specific epithet mamillana unfortunately has priority
over glaziouana, necessitating this new combination.
494
APPENDIX III
A new species of Stereocaulon
On two occasions between 1964 and 1971, a very strange
sterile lichen was collected on bare lave rocks, first by Yale
Dawson on Marchena (40905), and later by Larry Pike on San
Salvador (55240). These consisted of single, minute cup-shaped
gray squamules with upturned margins exposing a white lower
cortex. They were obviously juvenile basal squamules but they
were much more chondroid than those of any local Cladonia. In
June, 1976, Harry Sipman collected mature pseudopodetia, which
suggested that this was possibly a Stereocaulon or a Gymno-
derma. Study by Dr. Isao Yoshimura determined that it could not
be Gymnoderma, and that it produced marginal apothecia (no
spores were found, however) and rod-shaped pycnidiospores.
Morphologically it matched the Beringian Arctic Stereocaulon
wrightii Tuck. and S. apocalyptum Nyl., but whereas these
contain atranorine with stictic and lobaric acid, respectively,
the Galapagos material contains atranorine with norstictic acid.
Because cephalodia and spores were not found on the Sipman
collection, a special trip to Cerro Azul was made in 1984 to
collect enough material for the Lichenes Exsiccati COLO, with
good results. The new species forms masses of crowded pseudo-
podetia up to 10 mm high on the vertical faces of uncovered lava
tubes along the steep upper slopes of the mountain. Cephalodia
as well as small numbers of apothecia without well-developed
spores were found. The name of the new species commemorates the
type locality: Volcan Cerro Azul on Isabela Island.
The occurrence of an Equatorial vicariad species related to
two Beringian ones raises the question not only of dispersal but
of the time-frame over which evolution of these chemically-
diversified species took place. Dispersal seems most likely to
have been by birds. Chemical evolution has to have occurred
within the lifetime of the Islands, which is estimated from
between one and seven million years. Morphologically the three
taxa are indistinguishable.
Stereocaulon azulensis Yoshimura and Weber, sp. nov.
Thallus primarius concavus, orbicularis, usque ad 1 mm
diam, evanescens; pseudopodetia laciniato-ramosis, dorsiven-
tralibus, usque ad 10-15 mm longis, 0.8-2.0 mm latis, marginibus
irregulariter crenulatis, paginis dorsalibus corticatis, plus
495
minusve concavis, griseis, ultimo rimosis, paginis ventralibus
ecorticatis, albo-tomentosis, venosis, basin versus nigrescens,
K+ lutescentibus, P+ leniter lutescentibus, acido norstictico et
atranorin continens. Apothecia sessilibus, marginalibus, usque
ad 0.4-0.5 mm diam, lecidiinis; sporae desunt. Pycnidia margi-
nalibus et dorsalibus, ostiolis nigris 0.2-0.3 mm latis, pycno-
conidia 4.5-5.0 mu longis, linearibus, rectis interdum arcuatis.
Cephalodia atrobrunnea, verrucosa, super phyllocladia, Stigonema
continentibus.
Primary thallus concave, orbicular, up to 1 mm diam, disap-
pearing; pseudopodetia laciniately branched, dorsiventral, up to
10-15 mm long, 0.8-2.0 mm wide, the margins irregularly crenu-
late; dorsal surface corticate, more or less concave, gray,
cracked in age; ventral surface ecorticate, white-tomentose,
veined, blackening toward the base, K+ yellow; medulla K+ yel-
low, P+ weakly yellow; norstictic acid and atranorine present.
Apothecia sessile, marginal, up to 0.4-0.5 mm diam, lecidiine;
spores not found; pycnidia marginal and dorsal; ostiole black,
0.2-0.3 mm wide; pycnoconidia 4.5-5.0 mu long, linear, straight
or sometimes more or less curved. Cephalodia blackish-brown,
verrucose, on the phyllocladia, containing Stigonema.
HOLOTYPE: ECUADOR. Galdpagos Islands. Isla Isabela: Volcan
Cerro Azul, south slope above Iguana Cove, 700 m.s.m., on mossy
rock, bank of ravine, 17 Jan. 1984, Weber & Beck , Lich. Exs.
COLO 645 (COLO).
ADDITIONAL COLLECTIONS: Isla San Salvador: nearly barren, recent
a-a lava flow near James Bay, 27 April 1971, Pike 2607; Isabela:
Volcan Cerro Azul, 500 msm, 22 June 1976, Sipman 63507, 700 msm,
Sipman 63525.
LITERATURE CITED
Ahti, Teuvo. 1961. Taxonomic studies on reindeer lichens
(Cladonia, subgenus Cladina). Ann. Bot. Soc. "Vanamo" 32:1-160.
44 plates.
Arvidsson, Lars. 1982. A monograph of the lichen genus
Coccocarpia. Opera Bot. 67:1-96.
Awasthi, D. D. 1975. A monograph of the lichen genus
Dirinaria. Bibl. Lichenol. 2:1-108.
Brodo, Irwin M. 1984. The North American species of the
Lecanora subfusca group. Nova Hedw., Beih. 79: 63-285.
Darbishire, 0. V. 1935. The Templeton Crocker Expedition
of the Calif. Acad. of Sciences, 1932. No. 23. The Roccella-
fee EOC. Calit.. Acad, Sci. °21°285-294.
496
Dodge, Carroll W. 1935. Lichenes. In H. K. Svenson.
Plants of the Astor Expedition, 1930 (Galapagos and Cocos Is-
ands) 3) AMer aan DOCS a2 7 ee Ls
1936. Lichens of the Allan Hancock Ex-
pedition of 1934, collected by Wm. R. Taylor. Hancock Pacific
Expeditions 3(3):33-46.
Farlow, W. G.. 1902. ‘Lichens’,, ‘p. 83-89,''In:| B. Lo Ropine
son, Flora of the Galapagos Islands. Proc. Amer. Acad. Arts
SC oo.
Galloway, D. J. & P. M. Joergensen. 1975. Erioderma so-
rediatum, a new lichen from New Zealand. Lichenologist 7:139-
LA?
Gradstein, S. R., & William A. Weber. 1982. Bryogeography
of the Galapagos Islands. J. Hattori Bot. Lab. 52:127-152.
Hale, Mason E. 1984. The lichen line and high water
levels in a freshwater stream in Florida. The Bryologist 87:
261-265.
Hamann, Ole. 1986. The El Nifio influence on the Galapagos
vegetation. pp. 299-330 in Robinson & del Pino (eds.). El Nino
en las Islas Galdpagos. El evento de 1982-1983.
Hooker, Joseph Dalton. 1846. An enumeration of the plants
of the Galapagos Archipelago, with descriptions of those which
are new. Trans. Linn. Soc. 20:163-233.
Howell, John Thomas. 1932. Up under the Equator. Sierra
Club Bull) p.) 79-82.
Imshaug, Henry A. 1951. The lichen-forming species of the
genus Buellia occurring in the United States and Canada. Un-
published Ph.d. thesis, 217 pp. University Microfilms.
1955. The lichen genus Buellia in Cen-
tral America. The Bryologist 58:277-287.
1955a. The lichen genus Buellia in the
West Indies. Farlowia 4(4):473-512.
Magnusson, A. H. 1955. A catalogue of the Hawaiian li-
chens. Ark. f. Bot. (2) 3(no.10):1-402. 8 plates.
Malme, G. 0. 1927. Buelliae itineris Regnellianae primi.
Ark." tf) Boul 2 Aga 442.
Poelt, J., & A. Vezda. 1981. Bestimmungsschltissel euro-
pdischer Flechten. Erganzungsheft II. 390 pp. J. Cramer.
Redinger, Karl M. 1936. Thelotremaceae Brasilienses ...
Ark. (fi Bot. 265A 05): lel 22.,
Santesson, R. 1939. Uber die Zonationsverhadltnisse der
lacustrine Flechten einiger Seen in Anebodagebiet. Medd. Lunds
Univ. Limnotog wi inst . ls La7/0'.
Stewart, Alban. 1912. Notes on the lichens of the
Galapagos Islands. Proc. Calif. Acad. Sci. (4) 1:431-446.
Svenson, Henry K. 1935. Plants of the Astor Expedition,
1930 (Galapagos and Cocos Islands). Amer. J. Bot. 22:208-277.
497
1963. Opportunities for botanical study
on the Galapagos Islands. Occas. Pap. Calif. Acad. Sci.
44:53-58.
Tehlér, Anders. 1983. The genera Dirina and Roccellina
(Roccellaceae). Opera Bot. 70:1086.
Thomson, John W. 1963. The lichen genus Physcia in North
America. Nova Hedwigia, Beih. 7:1-172. 47 maps, 25 plates.
Usinger, Robert Leslie. 1972. Autobiography of an
entomologist. Mem. Pacific Coast Ent. Soc. Vol. 4. 330 pp.
California Academy of Sciences.
Weber, William A. 1966. Lichenology and bryology of the
Galapagos Islands, with check lists of the lichens and bryo-
phytes thus far reported. Pp. 190-200 [in] Robert I. Bowman
(ed.). The Galapagos. Univ. of California Press.
1971. Four new specles of Buellia from
western North and South America. The Bryologist 74:185-191.
1976. Additions to the bryophyte flora
of the Galapagos Islands. Lindbergia 4:76-79.
1981. Lichenes Exsiccati distributed by
the University of Colorado Museum, Boulder, Fascicles 1-15,
Nos. 1-600, 1961-1979. Mycotaxon 13:85-104.
1984. Galapagos cryptogams and the 1983
record El Nino. Internat. Lichen Newsletter 17:15-17.
1984a. Bryophytes and lichens of the
Galapagos Islands. National Geographic Society Research Reports
£Ot 29755716: /781-790-
, & Hans Beck. 1986. Effects on
cryptogamic vegetation (lichens, mosses and liverworts).
Chapter 3, in El Nifio en las Islas Galdpagos: El Evento de
1982-1983. 16 pages, 3 plates. Charles Darwin Foundation.
, & S. R. Gradstein. 1984. Lichens and
bryophytes. Chapter 5 [in] The Galdpagos Islands. Pergamon
Press, Key Environment Series.
, 9. R. Gradstein, J. Lanier & H. Sipman.
1977. Bryophytes and lichens of the Galdpagos. Noticias de
Galapagos 26:7-11.
(ewe Lanter, 't97). i Lichens! inthe
Galdpagos Islands. 20 pp. (in Spanish). Charles Darwin
Research Station.
Wiggins, Ira L., & Duncan M. Porter. 1971. Flora of the
Galapagos Islands. 998 pages, 96 plates, 268 figs. Stanford.
Willey, H. 1890. A synopsis of the genus Arthonia.
Privately printed.
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MYCOTAXON
Vol. XXVII, pp. 499-502 October-December 1986
PLATYGLOEA ACANTHOPHYSA, A NEW SPECIES WITH
SINGLE STERIGMATE BASIDIA AND ACANTHOPHYSES
HAROLD H. BURDSALL, JR.
Center for Forest Mycology Research
Forest Products Laboratory!
USDA, Forest Service
Madison, WI 53705-2398
The genus Platygloea Schroter in the Heterobasidio-
mycetidae is distinguished by a well-developed probasidium
that produces a cylindrical, transversely septate metaba-
sidium. The basidiospores germinate by repetition. How-
ever, several species of Platygloea form only an aseptate
metabasidium or a metabasidium with a single septum, e.g.,
Platygloea unispora Olive (1944) and P. mycophila
Burds. et Gilbn. (1974). Larsen (1984) indicated that
Laeticorticium simplicibasidium Linds. et Gilbn. (Lindsey
and Gilbertson 1977) is probably a single sterigmate member
of the Auriculariaceae as is Itersonilia perplexans Derx.
Two species of Septobasidium Pat. also possess
unisterigmate basidia, but they have a constant associa-
tion with scale insects.
The new species of Platygloea described here with an
aseptate metabasidium producing a single sterigma was
collected in northern Wisconsin in 1973. Further col-
lecting in similar habitats has not provided additional
specimens of this species.
1Maintained in cooperation with the University of
Wisconsin.
500
Platygloea acanthophysa Burds. sp. nov. Fig. 1
Basidiocarpo late effuso; hyphis fibulatis;
acanthophysibus variiformibus vel pyriformibus,
15-25 x 8-10(-13) wm, digitiformibus processibus usque
6 um longis; basidiis aseptatis, unisterigmatibus,
33-45 x 8-10(-13) wm; basidiosporis late ellipsoideis vel
subglobosis, 13-16 x 12-13 um, apiculis 2.5 x 2.5 wpm,
inamyloideis; acyanophilis.
Holotypus.--FP 101129, on Tsuga canadensis L., Kemp Bio-
logical Station, Minocqua, Oneida County, Wisconsin,
10 VIII 1973, leg. F. F. Lombard et R. G. Payne, in
CFMR conservatum.
Basidiocarp annual, broadly effuse, a thin vernicose
film, adherent, fertile area continuous, smooth to slightly
pubescent, pale gray; margin abrupt, not differentiated.
Hyphal system monomitic, subiculum an agglutinated
textura intricata, hyphae (Fig. 1C) 3-6 um diam.,
thin-walled or with slight wall thickening, hyaline,
smooth, with rare branching, nodose septate; subhymenium
not differentiated; acanthophyses (Fig. 1A) variously
swollen to pyriform, 15-25 x 8-10(-13) wm, hyaline,
thin-walled or with slight wall thickening, digitate
processes up to 6 um long, 0.5-1 wm diam., nodose septate
at base; basidia (Fig. 1B), including sterigmata, 33-45 um
long; probasidium poorly differentiated but usually with
slight wall thickening, 8-10(-13) wm wide, hyaline, smooth
or with acanthophysoid processes; basidiospores (Fig. 1D)
broadly ellipsoid to subglobose, 13-16 x 12-13 um, with
apiculus up to 2.5 x 2.5 um, hyaline, thin-walled, smooth,
nodose septate at base, not staining in Melzer's reagent,
acyanophilous, germinating by repetition.
Etymology.--From the acanthophyses present in the hymenium.
Discussion.--Although this species is known from only one
collection, it is so distinctive that it deserves to be
recognized. It is unique not only in the genus Platygloea
but throughout the Heterbasidiomycetidae in possessing
acanthophyses such as are found in the Aleurodiscoideae.
The digitate processes are also found on some basidia.
In addition to the acanthophyses, the single-spored
basidium and large broadly ellipsoid to subglobose
basidiospores are distinctive.
a
501
Fig. 1. Line drawings of microscopic structure of P.
acanthophysa. A. acanthophyses; B. unisterigmate
basidia; C. subicular hyphae; D. basidiospores, some
showing germination by repetition. (ML86 5208)
Platygloea acanthophysa, with basidiospores
12-18 x 9-14 um, is distinguished because of its pustulate
basidiocarp, the presence of simple or somewhat branched
hyphidia, and a long narrow basidium 61-83 x 6.1-7.6 um.
Laeticorticium simplicibasidium is distinguished from
P. unicornis in possessing globose basidiospores 9-11 wm
broad, basidia up to 125 x 3-5 wm. Itersonilia perplexans
Derx. has similar morphology but differs in possessing
basidiospores (conidia of Olive 1952) measuring
"7.6-10.6 x 12.2-18 yw" (Olive 1952). None of the species
mentioned previously possesses acanthophyses found in
P. unicornis.
The two one-spored Septobasidium species have a
constant association with both living trees and scale
insects, neither of which is associated with Platygloea
502
acanthophysa. Morphological differences between P.
acanthophysa and the Septobasidium species readily
distinguish them (Couch 1938).
The generic disposition of P. acanthophysa was not
straightforward. There was the possibility that it was a
one-spored species in the Tremellaceae. At the suggestion
of Dr. Kenneth Wells, University of California, Davis, CA,
and Dr. Franz Oberwinkler, University of Tubigen,
West Germany, I examined the septal apparatus by trans-
mission electron mycroscopy. The septa were distinctly
simple with no indication of a dolipore. This is the same
septal type present in the genus Platygloea. Species in
the Tremellaceae possess dolipore septa. Thus, the genus
Platygloea is the appropriate placement.
At the Forest Products Laboratory, Dr. M. J. Larsen
prepared the Latin diagnosis, Mr. T. A. Kuster performed
the transmission electron mycroscopy, and
Mrs. F. F. Lombard contributed a valuable manuscript
review. Dr. E. L. Stewart, University of Minnesota, and
Dr. Kenneth Wells also contributed valuable manuscript
reviews.
LITERATURE CITED
BURDSALL, H. H., Jr., and R. L. GILBERTSON. 1974. A new
species of Platygloea occurring on Peniophora tamaricicola
in Arizona. Mycologia 66:702-706. COUCH, J. N. 1938.
The genus Septobasidium. Univ. of North Carolina Press, |
Chapel Hill. 480 p. LARSEN, M. J. 1984. Notes on
laeticorticioid fungi. Mycologia 76:353-355.
LINDSEY, J. P., and R. L. GILBERTSON. 1977. New species
of corticioid fungi on quaking aspen. Mycotaxon 5:311-319.
OLIVE, L. S. 1944. New or rare heterobasidiomycetes from
North Carolina I. J. Elisha Mitchell Sci. Soc. 60:17.
OLIVE, L. S. 1952. Studies on the morphology and cytology
of Itersonilia perplexans Derx. Bull. Torrey Bot. Club
79:126-138.
MYCOTAXON
Vole XXV DI seppaets 03-5 06 October-December 1986
pak Ey URI SETA, ASE SESE ia al lea eh SEA oe aaa Sh Sa a Ns
MACROCHEMICAL COLOR REACTIONS OF MACROMYCETES
II. Chemical properties and systematic position
of Bondarzewia mesenterica
Harald M. Frank
Amthorstr. 5, Gera 6500, DDR
The discussion about the systematic position of Bondar-
zewia Sing. was renewed by Singer (1984) recently. Espe-
cially the similarity of the spore ornamentation has lead
to the assumption of common ancestry of Russulaceae and
Bondarzewiaceae. Newly recorded lactiferous species of
Bondarzewia and parasitic species of Lactarius Pers.
(Kreisel 1983, Singer 1984) are further reasons.
Carrying out systematic examinations of the macrochemical
behavior Bondarzewia mesenterica (Schaeff.)Kreisel (hit-
herto known as Bondarzewia montana (Quel.)Sing.) was
found to give a reaction pattern which seems to support
the close relation between these two families.
Only fresh specimen of single collections were tested.
The selection of the reagents, their concentration and
the use of the color atlas of Kiippers (1984) were founded
in detail by Frank (1987). Colors were regarded in each
case to be a composition of yellow (Y), magenta red (M)
and cyan blue (C). To simplify typing the format used
here is 30.10.70 which means Y 39M 9°79 or 30% saturation
in Y, 10% in M and 70% in C.
The abbreviations "n.ex." and "-" mean that the fruit
body was not examined and no change occured compared with
a piece exposed to air only. The nomenclature of Russula
Pers. is according to Kreisel (1983).
The reagents are KOH, H5SO0 4 = Hes, Feso0 , = FES, AgNO, =
AGN, phenol = PHE, l-naphthol = NAP, benzidine = BEN,
guaiac = GUA, guaiacol = GCL, ninhydrin = NIN, aniline =
ANI, formalin = FOR, Lugol = LUG and sulfovanillin = SV.
Russula caerulea (Pers.)Fr. emend. Cke. : KOH -, H2S8S -,
FES -, AGN -, PHE dark brown 55.85.80, NAP black violet
30.80.90, BEN bright greenish blue 25.00.99, GUA dark
blue green 30.30.99, GCL bright orange 70.50.00, NIN -,
ANNI se ROR Se LUG oe SV) air =.
504
Russula atropurpurea (Krbh.)Britz. : KOH yellow 30.03.00,
H2S greyish 05.10.20, FES greyish pink 10.30.10, AGN -,
PHE brown 35.50.40, NAP pale lilac 05.25.10, BEN olive
80.20.40, GUA bright blue 25.10.90, GCL apricot 35.25.00,
NIN -, ANI -, FOR -, LUG yellow brown 70.50.40, SV violet
10.60.65, air -.
Russula chloroides Krbh. : KOH yellowish 20.00.00, H2S -,
FES rapidly pink 15.30.00, AGN -, PHE vinaceous to dark
brown 70.80.60, NAP blackish violet 20.80.60, BEN in-
stantly bright blue 10.00.80, GUA very rapidly dark blue
green 30.20.80, GCL red brown 60.60.30, NIN slowly
pinkish 10.20.00, ANI -, FOR -, LUG greenish brown
COTSONIO COV Telex a terra
Russula fellea (Fr.)Fr. : KOH strong yellow 60.00.00, H2S
dun 40.40.60, FES pale greyish pink 10.10.05, AGN strong
chocolate 50.80.80, PHE red brown 60.60.40, NAP -, BEN
ochraceous brown 35.50.40, GUA dull dark green 50.35.85,
GCL red brown 60.60.40, NIN -, ANI -, FOR -, LUG yellow-
ish brown 90.65.65, SV purple 25.75.55, air <-.
Russula ochroleuca (Pers.)Fr. : KOH yellowish 20.00.00,
H2S -, FES greyish pink 10.20.05, AGN pale pink 10.10.00,
PHE dark brown 50.70.80, NAP dark violet 30.70.80, BEN
instantly light blue 10.00.65, GUA instantly blue green
30.00.75, GCL bright red brown 70.80.60, NIN -, ANI pink
greyi30.20°520,° FOR pale) pank).10.10,00,,..LUG <—, SVrm.e2..
air mast |
Russula virescens (Schaeff.)Fr. : KOH yellowish 20.00.00,
H2S pale yellowish 10.00.00, FES rather instantly pink
15.30.00, AGN -, PHE greyish pink 15.25.03, NAP greyish
lilac 05.20.05, BEN apricot 50.10.00, GUA rapidly dark
blue green 30.30.99, GCL pinkish orange 25.40.00, NIN
very pale grey 05.05.05, ANI yellowish 20.00.00, FOR -,
LUG bright yellow 70.05.05, SV bright purple 03.50.03,
air =e
Russula cyanoxanthe (Schaeff.)Fr. : KOH -, H2S -, FES
greyish 10.05.10, AGN pale pinkish grey 10.10.05, PHE
vinaceous brown 70.85.60, NAP violet 10.45.45, BEN in-
stantly bright blue 10.00.99, GUA instantly bright blue
25.00.99, GCL vinaceous lilac 25.50.20, NIN very pale
bluish violet 05.10.20, ANI ochraceous pink 35.25.00,
FOR -, LUG -, SV n.ex., air -.
Russulea heterophylla (Fr.)Fr. : KOH yellowish 20.00.00,
H2S -, FES instantly reddish 15.30.00, AGN yellowish
creme 30.10.00, PHE rapidly red brown 70.70.50, NAP rapid-
ly violet 10.45.45, BEN instantly blue 10.30.80, GUA in-
stantly dark blue green 30.30.99, GCL dull yellow
25.10.05, NIN slowly grey 10.05.30, ANI very pale greenish
yellow 10.00.05, FOR n.ex., LUG -, SV n.ex., air -.
ous
Russula xerampelina (Schaeff.)Fr. : KOH yellow 50.00.00,
H2S -, FES greyish green 30.10.40, AGN greyish pink
10.30.10, PHE dark red brown 70.80.60, NAP dark violet
30.70.80, BEN instantly bright blue 10.00.90, GUA rapidly
dark green 50.35.85, GCL red 15.55.05, NIN greyish pink
10.20.00, ANI bright red 45.75.05, FOR greyish pink
10.20.00, LUG -, SV -, air -.
Bondarzewia mesenterica : KOH golden 55.25.00, H2S grey
20.10.20, FES -, AGN -, PHE greyish 10.10.10, NAP rapidly
violet 00.50.50, BEN instantly bright blue 10.00.99, GUA
rapidly dark blue green 30.30.99, GCL rapidly bright
orange red 70.60.00, NIN -, FOR -, LUG instantly blackish
brown 80.90.90, SV dark violet 30.70.80, air -.
The species of Russulaceae are very profitable objects
for agaricologists due to their various behavior against
different reagents. The strong discoloration after stres-
sing with oxidase-sensitive molecules like phenol, ben-
Zidine, l-naphthol, guaiac and guaiacol is a rather
unique phenomenon comparable at the present stage of
knowledge only with species of the genus Agaricus L.
In a representative 72 specimen sample Marr (1979) found
that the combination of laccase and tyrosinase is a typi-
cal and constant feature of the 13 Russulacese species
examined. Additionally these specimen represent about
90% of the specimen with the coupled oxidases.
Therefore it is interesting to note that Bondarzewia me-
senterica is very similar to the majority of the Russula
species choosen here. The white fruit body contains lar-
ger amounts of different oxidases and reacts with sulfo-
vanillin. The latter is according to Camazine and Lupo
(1984) probably an indicator of the pungent principle
(acrid tasting aldehydes) common and typical in Russula-
ceae. The compounds detected here are not only secondary
metabolites but reflect as well physiological potency as
ecological requirements.
It seems to be justifiable to plead for the concept of
Singer (1984).
Differences within the genus Russula e.g. the ochraceous
color of Russula fellea after applying benzidine (an ex-
traordinary reaction which was already described by Micka
and Klan 1980) should be understood as a result of phylo-
genetic divergence.
The results support the authors intention in 2 respects:
macrochemical color reactions are usefull to distinguish
closely related species (e.g. Russula fellea and Russula
ochroleuca) and/or they can be used as arguments in
establishing phylogenetic pathways.
Acknowledgement
I wish to thank R. Conrad and F. Groger for their help
in collecting and examining material.
Literature
Camazine, S. & A.T. Lupo, Jr. 1984. Labile toxis com-
pounds of the Lactarii: The role of the laticiferous
hyphae as a storage depot for precursors of pungent
dialdehydes. Mycologia 76(2): 355-358.
Frank, H. L987. Makrochemische Farbreaktionen bei
GroBpilzen I. Voraussetzungen fiir eine systematische
Untersuchung. Z. Mykol., accepted for publication.
Kreisel, H. 1983. Michael-Hennig-Kreisel, Handbuch
fiir Pilzfreunde, Bd. 5. Fischer Verlag, Jena. 408 p.
Kiippers, H. 1984. DuMont's Farbenatlas. DuMont Buch-
verlag, Koln. 164 p.
Merr, C.D. IBS Wi SIe Laccase and tyrosinase oxidation of
spot test reagents. Mycotaxon 9: 244-276.
Micka, K. & Jd. Klan. 1.980% Chemical spot tests of
macromycetes with benzidine. Ceska mycologie 34(2):
74-81 °
Singer. R. 1984. Tropical Russulaceae II. Lactarius
section Panuoidei. Nova Hedwigia 40: 435-442.
MYCOTAXON
VOU NAVI Pp.) 07> 550 October-December 1986
cS SR EE a EL el al ce a eat Ci ah ak ed Ne a
EAST AFRICAN RUSTS (UREDINALES), MAINLY FROM UGANDA
5. ON FAMILIES BELONGING TO GAMOPETALAE
Halvor B. Gjaerum
Norwegian Plant Protection Institute
P.O. Bax 70, N-1432 AS-NLH, Norway
Summary
seventy-one rust taxa on host families belonging to
Gamopetalae (Sympetalae) Of: .the Dicotyledones are
recorded. Ten species are described as new, viz. Aecidium
aspiliae on, Aspilia’ pluriseta, A. echuyaénse on
Helichrysum foetidum, A. haplocarphae on Haplocarpha
rueppellii, Puccinia artemisiae-afrae on Artemisia afra,
Ee, crassocephali on Crassocephalum aframontanum Pe
tetradeniae on Tetradenia riparia, Uredo hypolstis-
verticillaris on Hypo€stes verticillaris, U. isoglossae on
Isoglossa substrobilina, Us leucadicola on Leucas
martinicensis, Uromyces sasalnsis on Valeriana volkensii.
Uredo paederiae on Paederia foetens is new to Africa.
Eighteen species are reported as new to Uganda, three are
new to Kenya, and one to Tanzania.
The main part of material treated was collected by
myself in Uganda in 1970 or picked out, from herbaria in
Kampala and Oslo. For comparison I have borrowed material
from Swedish Museum of National History (S), Royal Botanic
Gardens, Kew (K), and Commonwealth Mycological Institute
(IMI). My thanks are due to the curators of the herbaria
mentioned.
During my stay at CMI in 1982, kindly financed by the
Norwegian Agency for International Development (NORAD), I
alco menad the. Opportunity, toapuse. their’ material tor
comparison, and for which I am most grateful.
Me, Uh wan buuVvermat othe ywotanical si Insticure.” of ythe
Agricultural University of Norway has kindly helped me
with, host “identifications and Dr. G.By’ Cummins, Dept. of
Plant Pathology, University of Arizona, Tucson, AZ, and
Prof. D.M. Henderson, Royal Botanic Garden, Edinburgh,
have critically read the manuscript. I am much indebted
for this valuable help.
Finally, sl eiwant “to ithanky. heartily - Mrs. “Rannveig
@degaard, Norwegian Plant Protection Institute, for typing
508
and correcting these manuscripts dealing with East African
rust cunga.
Aecidium aspiliae n.sp. (Fig. 1)
Pycnia epiphylla in maculis flavis, typus 4. Aecia
hypophylla, in gregibus parvis aggregata, peridio albo,
lacerato, revoluto. Cellulae peridii rhomboideae, 30 x 20
pm, pariete interno verrucoso, 3 pm crasso, pariete exter-
no \striato, 9 gm crasso. Aeciosporae oblongae vel angu-
lares, ‘6-20 ' x7 13=15 pn, pariete 0.5 pm crasso, hyalino,
verruculoso, saepe particulas paucas parvas’ lucentes
continentes, poris obscuris.
Holotype: | H.B. Gjarum 71 (MHU), 09.09.1970, Uganda, W.
Mengo, Busiro Co., Zika Forest, 1160 m, on Aspilia pluri-
seta Schweinf. (Asteraceae). Isotype in NPPI.
Several rust species are described on the genus
Aspilia, but the only one having aecia is Puccinia fragi-
lispora Dale, described from Trinidad. According to the
diagnosis (Dale 1955) it has much larger peridial cells
and aeciospores, 30-70 x 20-41 and 30-48 x 20-27. pm,
respectively.
enhean Sy
- *, s "tn, 82M, aie M02 ¢0N0 gph,
§ % 6g 3 mE! 3
§ / & : 3 :
Wo) ae ane 6 A Soins ia aie
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el) ‘Ga ea Cat? bs iTGoy % ‘i a
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a. (8) Ge! Pe ed
Fig. 1. Aecidium aspiliae n.sp.
Aeciospores from type.
Aecidium;cynanchiCcumm.:, | Bull. Torrey Bot...) (Club veo ssa,
1941.
On Cynanchum cf. altiscandens K. Schum. (Asclepiadaceae)
Kenya. Central Prov., Suam Forest Station, 2100 m, 23.-
24.0129 AS ieeevvanrden 69276 Ope.
No pycnia seen. Aecia hypophyllous in yellow leaf
spots. Peridium white, tubeformed, slightly lacerate and
revolute at the top. Aeciospores irregularly angular, 15-
21 x 13-18 pm, wall 1 wm thick, hyaline, verruculose.
This species which is new to the rust flora of Kenya,
was described from Sierra Leone on C. mannii N.E. Br. ,
and it is later reported on the same host from Ghana. C.
altiscandens is a new host for this rust.
509
Aecidium dichrocephalae P. Henn., Monsunia 1: 4, 1899.
On Dichrocephala integrifolia O. Kuntze (Asteraceae)
Bugisu Distr., N. Bugisu Co., Bumasifwa, above the school,
1200tme Gs. (404) Ankole Distr, Bunvaruguru. Co. ,\Kalinzu
Forest, 11400 M,.f G7.) (i403 KRiges, Distr.’, _Bufumbira \Co.,
on track to Gahinga, just above the border of cultivated
LANG 2oUOUTn, Gaia (i241).
Pycnia epiphyllous, yellow brown. Aecia hypophyllous,
in great clusters. Peridium white, revolute, lacerate.
Aeciospores ellipsoid, ovoid or irregular, 25-33 x 15-20
Am, wall 1 pm thick, at apex thickened up to 8 pm, hyaline
and densely verrucose.
This aecial stage has been reported by Wakefield &
Hanstora (1949) on Di \ latifolia) DC. from Kigez1i ‘and .on D.
sp. from Bugisu, and by Ebbels (1972, as A. matapense
Cumm., IMI 155315) from Kigezi. Outside Africa it is known
on D. spp. from Java, Assam, Taiwan, and New Guinea.
Aecidium dubiosum H. & P. Syd., Hedwigia 40: (1), 1901.
On Senecio sp. (Asteraceae)
Bugveuybi ser, ; iN. \ouerstiCo., Mt. bigon, ab. Sasa, River on
track tosasa Hut, 2650 m, Gp. (386) 5:0 4 2.
Pycnia epiphyllous, yellow-brown. Aecia hypophyllous
in great clusters. Peridium white, revolute, lacerate.
Aeciospores ellipsoid, ovoid or irregular, 24-33 x 15-21
pm, wall 1 pm thick, at apex thickened up to 7 pm, _hya-
line, densely verruculose, pores obscure.
A. dubiosum, a new rust species in Uganda, has been
reported on several Senecio spp. in Africa, esp. from S.
Africa, but it is also known in Malawi. Outside Africa it
has been found in Portugal.
Aecidium echuya€nse n.sp. (Fig. 2)
Pycnia epiphylla, diam 130 um, typus 4. Aecia hypo-
phylla, in gregibus parvis aggregata, peridio albo, longo.
Cellulae peridii usque 40 pm longa, pariete 4 pm crasso,
pariete interno striato, pariete externo verruculoso.
Aeciosporae ellipsoideae vel subgloboideae, 23-33 x 18-27
pm, pariete 1 pm crasso, hyalino, verruculoso, poris
obscuris.
HOLOCy pean eoe wGlerul 230A rUMHU Ol. bOstorvO,. “Uganda,
Kigezi Distr., Rubanda Co., Echuya Forest Reserve, Mile
34-35 on the Kabale-Kisoro road, 2100 m, on Helichrysum
foetidum (L.) Cass. s. lat. (Asteraceae). Isotype in NPPI.
In the literature available to me three aecial stages
are mentioned on Helichrysum, two of which are described
with apurtenant telial stages, viz. Puccinia macowanii
Wint. and P. pinarii Pole Evans. Doidge (1926) when
describing Aecidium helichrysi indicated that it might
belong to P. kalchbrenneri De Toni. The latter has thicker
spore wall, 2.5-3 pm and P. macowanii has thicker peridial
cell walls and more globose spores than described for the
new species. The aecial stage of P. pinarii might be that
which is closest but the spores have a granular content
510
which the spores of A. echuya@nsis do not have.
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Aecidium echuya¥nsis Nesp.
Aeciospores from type.
Aecidium*flavidum Berk, & Bre, dle’ Linn. Sec. Londen via:
OS pu B72
On Pavetta schumanniana K. Schum. (Rubiaceae)
Tanzania. Iringa Distr., Ruaha National Park, at Magangwe
AIT Strip, 1340sm, 11.03. 19/27 A. Bjarnstad, C1540) (0) ae.
Pycnia epiphylla. Aecia hypophyllous, in groups. on
yellow-brown leaf spots. Peridium brownish (old), revo-
lute. Spores subgloboid or angular, 23-32 x 20-23 pm, wall
T=155 pm thick, verruculose and» with refracting granules.
A. flavidum, widespread in Tropical Africa and Asia,
was reported from Uganda by Wakefield (1920) and Wakefield
& Hansford (1949) on Pavetta sp. from Kivuvu in Kyagwe.
The host is new to the fungus in Uganda, but has_ been
reported from Zambia.
Aecidium gynurae Petch, Annal. R. Bot. Gdns Peradeniya 5:
XW OAM SER
Syn. A, sformosanum syd.) Annis imycol. clits (5Oy 071s aee es
emiliae "Petchy Annal?’ R.” Bot. (Gdns Peradeniya 6:9 2127
OT seers. nuree March, . & Stayert. Bull. Soc 3 yr @eoc.
Belg: °61,° "sere" 227 165, 19193 A. scrassochephalatWakerees
Hansf ..*-Proc.- Linn. -Soc; Lond.,; Sess. 161. 1948-49 (Pu.
189, 1949; A. gynurae-cernuae Vien.-Bourg., Bull. Soc.
Mycol, VFiroe poe eat i | 955.
On Crassocephalum crepidioides (Benth.) DC. (syn. Gynura
crepidioides Benth.) (Asteraceae)
W. “Mengo <Distr., - Kyadondo Coy, Makerere Hild) 12007 em,
Taksdale sc 1Sye (Gye Dy16. ht 1970 (flegses..Ccusital lant ies
Kabanyolo, 1200 m, Gj. (487); Mawakota Co., Mpanga Forest
1150 m5 Gis 7 792 Busito ‘Co., "Kisibawoe1200cmypaGi. ec,
Entebbe “1150°m, Gis -C38)4 VES “Mengo Distrs., Kyagweuco.,
near Bugombe on Kome Island, 1150 m, 27.10.1968, K.A. Lye;
Teso Distr., Serere Co., Serere Research Station 1150 n,
Cyr C431) er Oray bh.
On C. rubens (Jacq.) S. Moore
Mupendel Dest, ~Singo- Co.," Homme, 1500 em," Gy. 8¢325 ys
Bunyoro” *Distr., Buruli Co’, “Bunyoro Ranch near Masindz
20ehe6. 1961, Turner (452:);,°°O 4a.
Pycnia epiphyllous, type 4, rare. Aecia in dense
clusters, mainly on the lower side of circular, yellow-
brown leaf spots. Peridium short, revolute, white to
yellowish. Aeciospores angularly globoid or ellipsoid, 16-
19 x 14-16 ppm, wall ijpm? thick, “hyaline, verruculose anc
with some refracting granules.
To my knowledge pycnia have not been mentioned
earlier, and they occur in most specimens examined.
This aecial stage is widespread in tropical Africa
and in Asia east to the Philippines, Taiwan and the
Ryukyus, and it is also reported from Queensland. The main
host genus in Africa is Crassocephalum (incl. Gynura). In
Asia Emilia is the main host genus, but Gynura and Senecio
are also reported.
Viennot-Bourgin (1958) recognized three Aecidium
species, A. gynurae Petch (syn. A. emiliae and A. formosa-
num), A. mariani-raciborskii Siem. andA. crassocephali
(syn. A. gynurae March. & Stayert and A. gynurae-cernuae
Vien.-Bourg.) while Bouriquet & Bassino (1965) placed
them within A. crassocephali. After having studied the
Ugandan material presented above and the types of A.
gynurae Petch and A. crassocephali (Herb. K) as well as
their diagnosis, I find it most convenient to treat them
aS one species. According to the diagnosis, the third
species recognized by Viennot-Bourgin, viz. A. mariani-
raciborskii, has larger spores than have the other species
discussed. As the type specimen probably has been lost (T.
Majewski in litt.) and the spore measurements thus cannot
be verified, I find it better to keep it separate.
Wakefield & Hansford described A. crassocephali on C.
crepidioides from Busoga. C. rubens is a new host for this
rust in Uganda, but it has been reported as a host from
Tanzania and Malawi.
Aecidium haplocarphae sp. nov. (Fig. 3)
Pycnia non visa. Aecia amphigena, pro maximam partem
hypophylla, in maculis ferrugineis plus minusve rotunda-
tis, diam 300-400 fem, aggregata, peridio lacerato, revo-
luto, juvenilio albo, demum pallide brunneo. Cellulae
peridii rhombohydrales, imbricata, pariete externo 4 pm
Crassoy, “Striato, ~pariete” interno” 2 ppm crasso, gYrosso
verruculoso. Aeciosporae globoideae, ovoideae vel poly-
hedroideae, 17-24 x 14-18 pm, pariete pallide brunneo,
1.5=-2.5 pm crasso, dense verruculoso, saepe particulas
paucas parvas lucentes continentibus, poris obscuris.
Holotype. fn. B. Giercun 412" “(MHU)S °2451021970, “Uganda,
BUGtSW@oDrsth., sh. BuGaSsu CO,., Mt. Blgon, between Sasa Hut
and Jackson's Summit, 3800 m, on Haplocarpha rueppellii
(Sch. Bip.) Beauv. (Asteraceae). Isotype in NPPI.
The host occurs at an altitude between 2750 and 4700
m in the mountains in Ethiopia, Kenya, Uganda and
Suz
Tanzania. /At-Mt. | Elgon it was found /up.to4300 .m.sSottan
no rust seems to be reported from this host. Nattrass
(1961) reported a Uredo sp. on the closely related H.
schimperi (Sch. Bip.) Beauv. from Kenya, but the specimen
in CMI (IMI 45433) is recorded as a species of Landtia.
Fig. 3. Aecidium haplocarphae n.sp.
Aeciospores from type.
Aecidium macowanianum Thuem., Flora 58, 380, 1875.
On Conyza sp. (Asteraceae)
Kenya. Eastern Prov., Nyeri Distr., Mt. Kenya, S slope,
Regati- Forest Station, 2200-2300 m;)) 02.-03:.02.V973, seiae
Ryvarcen 109112 300) abe
Pycnia not seen. Aecia amphigenous, in dense clus-
ters. Peridium white, revolute, lacerate. Aeciospores
subgloboid or somewhat angular, 15-21 x 13-19 pm, wall 1
fam thick, in some spores thickened to 2,5 pm at apex,
hyaline, verrucose.
This aecidial stage was reported by Wakefield &
Hansford (1949) .0on,C. hochstetteri Sch. Bip.sex A. Rich.
from Ankole and on C. sp. from Kabale in Kigezi and from
Kampala. On other Conyza spp. it has been reported from
Kenya, Tanzania, Angola. and S.* Africa, andsalso from
India.
Henderson (1972) compared A. macowanianum with A.
conyzae-pinnatilobatae P. & H. Syd. The latter is said to
be thickened at the apex, while the former has_ evenly
thick-walled spores. In his material Henderson found that
it showed "slight signs of irregular wall thickening so
the two species may not be absolutely distinct". The Mt.
Kenya specimen shows in some spores ae slight wall
thickening, but not as much as given in the diagnosis for
A. conyzae-pinnatilobatae (in the diagnosis up to 5 pm).
Another species, A. hoffmannii . Pw) Hees yeied
described.on Conyza limosa O. Hoffm. from S. Africa has
somewhat larger aeciospores with apex thickened up to 8
pm.
Aecidium, plectranthi ‘Barcl.,\,Jd, Asiat.,.Soc. Bend. S59eULi):
104, 1890.
Di
On Plectranthus sylvestris Glirke (Lamiaceae)
hagezi” Distr... Botumbira” {Co., on. Jsthe, saddle” between
Muhavura and Gahinga, ~3]00)m, Gj. (256), .1.
No pycnia seen. Aecia amphigenous, scattered or in
small groups. Peridium white, tubeformed, slightly lace-
tavevand revolute, cells 39-55% 19-23) pm, walls) 3-3. 5) jam
thick, outer wall striate, inner wall verrucose. Aecio-
spores obovoid to ellipsoid, 27-32 x 18-27 pm, wall 1.5 pm
thick, hyaline, verrucose, pores obscure.
In addition to E. plectranthi two more species with
aecia are described on Plectranthus, viz. A. plectranthi-
cola .Cumm. and Puccinia plectranthi Thuem. The former
differs from A. plectranthi having aeciospores' strongly
thickened at the apex. The aecia of P. plectranthi were
poorly described by Cooke (1882), only said to be hypo-
phyllous with yellow spores. The specimen studied by me
lacks pycnia and has amphigenous, mainly epiphyllous
aecia. Spore contents pale yellowish. As there are no
telia I prefer to place it with A. plectranthi (see also
Dordge 1926).
A. plectranthi is new 'to the rust flora of Uganda. In
AteLeca se iitaunis gleponted on Pi. flomntbundus, N.BsBr.. uhtoom
sanbabwe and 5. Africa,..on Pl. sp. also from Tanzania. /In
Asia it is known from N.W. Himalaya, China, Japan and E.
Siberia. P. sylvestris is a new host for this rust.
Aecidium pycnostachydis (Kalchbr.) Doidge, Bothalia 2:
(Eee OM ROPES
PV) Uredo —pyecnostachnydis Kalchbr.,, Grevillea 11t:) 25,
1882.
On Pycnostachys meyeri GlUrke (Lamiaceae)
KigeciapiSstr., ) butnumbirarco., “inthe Tower ‘forest: border
on Crack cto: Gahinga, .2700 im, Gj}. (244), 1.
Pycnia not seen. Aecia amphigenous, scattered or in
small groups on brown leaf spots, cupulate. Peridium
white, revolute, lacerate. Spores angular to ellipsoid,
25-30 x 20-23 pm, wall 1.5 pm thick, verrucose with irre-
gular warts.
This aecial stage, described from S. Africa, was
reported by Wakefield (1920) and later by Wakefield &
Hansrord (1949) one. dawei N.E. » Br., ..from.Mugomba, in
Kyagwe. It has also been recorded from Zimbabwe. P. meyeri
as ainew host’ for this rust:
Aecidium vernoniae-podocomae Doidge, Bothalia 2: 212,
Mie di
On Vernonia auriculifera Hiern (Asteraceae)
W. Mengo Distr., Kyadondo Co., Namulonge and Kabanyolo,
200m, | 7). CS2°e2S0 re BUST YO oCO., Kiisibawoisl.20 Op im) sae
(190), Mawokota Co., Mpanga Forest, 1150 m, Gj. (174), all
ilies
Ethiopia. Shoa Distr., Wondo Gennet, 1800 m, Gj., I.
Aecia hypophyllous, yellow-brown, round, circular,
scattered or a few loosely together. Peridium white, often
514
broken. Aeciospores angular, 17-28 x 17-20 um, wall 1 pm
thick, hyaline, densely verruculose, with very small
refracting granules.
Wakefield & Hansford (1949) reported this aecial
stage on Vernonia sp. from Kawanda in Kyadondo.
Endophyllum (?) striatosporum Wakef. in Wakefield &
Hansford, “Proc. binn. Soc. Lond:, Sess." 16). 1948-497 pr
22 162-198) 1:949%
On Rhamnus prinoides L'Hérit. (Rhamnaceae)
Etnhropial choa Distr, >. yardgou 25S CIMl 165.07. ol oe
On Rhamnus sp.
Tanzania. Arusha Prov., Mt. Meru, E slope, Meru Crater,
2000 1m:,/?17 052970, (Ni Ls hiundaqvast (6502) 0UPS)=
Aecia densely clustered on leaves and hypertrophied
twigs, ‘peridium long, tubular, revoluted, ‘white- Spore—
masses yellowish. Aeciospores subgloboid to ellipsoid,
angular 1/-27 x 15220 pum, “wall 2.5-5 um thick, hyaline,
striate.
The rust was described on R. prinoides from Sabei in
Uganda. It has later been reported as E. macowanii Pole-
Evans on the same host from Tanzania (Henderson 1970) and
Ethiopia (Stewart “& “Yirgou 1967). "E. ‘macowanii,. alse
reported from Tanzania and other African countries from S.
Africa to Sudan, has verruculose spores.
Wakefield pointed out that she had not seen the spore
germination, but she placed the rust tentatively in the
genus Endophyllum "because of its suggestive resemblance
to E. MacOwani Pole Evans". To my knowledge nobody has
studied germination.
Endophyllum superficiale (Karst. & Roum.) Stev. &
Mandiola,; PRilipp Agric. -202 13 1931.
syn. Aecidium’clerodendri P.. ‘Henn: ; Bot. Jb. 15: 6,7) 1692.
On Clerodendron cordifolium (Hochst. ) A. Rich,
(Verbenaceae )
Acholi Distr.;,, “bamwoCo., 3) kmuSW of Lotuturi, “160GR. a,
17.02.1969, K.A. Lye & R.N. Lester (2040).
Sori round or irregular, thickened, bullate. Spores
oblong or rectangular 23-28 x 20-23 pm, wall 1 pm_ thick,
verrucose, hyaline.
The rust is new to Uganda, but it has been found in
Malawi and Zambia. In Asia it has been found from India to
China and Japan on several Asiatic Clerodendron spp. and
also on Sterculia in Vietnam. C. cordifolium is a new host
for the xsust.
Hemi leratcanthiiBerk. (6 Br.) J .-Linns Soc. Bots Gond.sue.
93h) MO toe
On Canthium quenzii Sond. (Rubiaceae)
Kenya. Kwale Distr., Shimba Forest, near Pengo Hill,
09.01.1970, A.B. Katende & K.A. Lye (4810), II.
Uredinia hypophyllous in small tufts, yellow when
er
35
fresh, when dry white. Urediniospores irregular, angular,
ellipsoid, often flattened at one side, depending on the
Cpmentationw Of the spore; e1c0=34 (3720-25e0m, vwall ~ 1° om
thick, verrucose, but partly smooth, hyaline.
Eevcanthit iastnew.co the ruse flora, of! Uganda,.and Cc.
Ciena S| omenew wnOSt Tora ten kin Arricay at. has) been
reported from S. Africa, Ghana, Angola and Tanzania on
Canthium ‘spp. and also on Plectronia sp. ‘In Asia it’ is
found in Pakistan, India, Sri Lanka and the Philippines on
the same host genera as in Africa, in Pakistan also on
Holarrhena antidysenterica Wall.
Heminerva molstii P.41ék HH. Syd., Monogr. Ured. 3%. 213, 1915.
One Psycnocria kirkis Hiern’ var. -hirtella’ (Oliv), Verdc.
(Rubiaceae)
Masaka Distr., BukOto. (Co-.); near Bugonzi, 1160 om.
30 ero 1), Ke Aau Lye, (M,) 56.B). Tt.
Uredinia amphigenous, mainly hypophyllous, small,
scattered or confluent, pulverulent. Spores obovoid to
globoid, flattened at one side, depended on orientation,
Zena ex No-20snm,) wallot pm thick, hyaline; roughly verru-
cose, but partly nearly smooth.
This rust has been found in Kenya, Tanzania, Zambia,
Malawi and Uganda, in the latter country on P. nigropunc-
Paves andihoayasp. \ (Waketteldaé: Hansford. 1949)... Po kirkii
Verunurcellacis, a vnew host for this cust.
Henmvercm ccno Uz iar h.. & P.oya.,) Bota. wb. 4557 260.191 0%
Suir credouscnouzi ys) Po) Henn Bob Jb. 333034, 1902.
On Clerodendron rotundifolium Oliv. (Verbenaceae)
Kigezi Distr., Rubanda Co., Mile 26 on the Kabale-Kisoro
MOcG eC OU, Gi. (210), 0s Bunyoro, Distr... Buyenje Co.,
Louewe sborenole,. 1050m, Gi). (335), 11.
Uredinia hypophyllous, in dense groups, orange-
yellow. Urediniospores 23-29 x 20-23 pm, bilaterally
ovate, lower part smooth or nearly so, wall hyaline, 1.5-
2i(=2 .5;) jan thick, echinulate.
Wakefield & Hansford (1949) reported this rust C.
yOnnstonii Oliv., C. capitatum Schum. & Thonn. and C. sp;
from several localities from the Kampala-Entebbe area _ to
Kigezi. -it. “is widespread “in:Africa from Sudan to 5.
Africa, and also known in Mauritius and Samoa.
C. rotundifolium is a new host for the rust.
Hemiteiva, vastatrix Berk. 4) Br.;, Gdns: Chron. 1669+: 1157.
On Coffea arabica L. (Rubiaceae)
W. Mengo Distr., Kyadondo Co., Kawanda, 1180 m, Gj. (491),
Tis) Bugisu Distr.) )Nw,;Bugisu Co., above Bumasobo school,
POU aaNet S50 ),. eller loro Distr... Busongora Cos. Near
Kyakikere on Zaire border, 1400 m, 21.04.1970, K.A. Lye
CM 1604 CL.
On C. canephora Pierre ex Froehner
W. Mengo Distr., Kyadondo Co., Makerere, 1200 m, Gj.
C456), (OSLO9ETS 70 (N.C. Lwanga,) ily Rabanyolo,, t20Griercae
(286), LR) Mawokota’ Co., Mpanga Forestii.1.150) mpsGale haven
LiL,
Uredinia hypophyllous, yellow-orange. Urediniospores
21-38 x 15-23 pm, wall nearly hyaline, roughly echinulate
and thicker on the upper (convex) side than on the lower
Side which is nearly smooth.
According to Wakefield & Hansford (1949) this rust is
common throughout the country. They?) also “report mes
liberica Hiern and C. excelsa Cheval. as hosts. Henderson
(1970) reported it on Gardenia jovis-tonantis Hiern from
Kasambya. The coffee rust is widespread in Africa, in
tropical Asia and Oceania, andalso in S. and C. America
from Argentina to Mexico.
Hemileia woodii Kalchbr. & Cooke, Grevillea 9: 22, 1880.
On Vangueria acutiloba Robyns (Rubiaceae)
W. Mengo Distr., Busiro Co., Kisibawo, near Mile 13 on the
Kampala-Hoima road, 1200 m, Gj. (186), “Li; Teso, Distua,
Serere Co., Serere Research Station, 1150 m, Gj. (449),
LER
Uredinia hypophyllous, sori small, in groups on
greyish brown leaf spots. Urediniospores bilaterally
ovoid, 27-34 x 14-20 pm, depending on the orientation of
the spore, wall 1.5 pm thick, “hyaline, echinulate’ except
for the concave side which is smooth.
This rust species, new to Uganda, is widespread from
Ethiopia through Kenya, Tanzania, Zambia and Zimbabwe to
S.’ JAfrica, “and it) is) also. known)/in india.) the) main nose
genus is Vangueria. V. acutilobia has been reported from
Tanzania.
Phakopsora)'vernoniae Gerst, ,/ “Ark. " Bot.) Ser. 2 esa.
SOs) a Or
On Vernonia amygdalina Del. (Asteraceae)
Wi.) iMengo Distr.) | Kyadondo Cosi, ‘Kabanyolo,’/ (1200) myiaa.
[270 ye ase
On Vernonia campanea S. Moore
W. Mengo Distr., Mawokota Co., Mpanga Forest, 1150 m, Gj.
C179) 5) Tas
On Vernonia juglandis Oliv. & Hiern
Kenya. Central Prov., Mt. Elgon, Suam Forest Station, 2100
mM, 238-24 0001973, 1b. Ryvarden: (9268) 0) pr leiite
Uredinia hypophyllous in brown leaf spots, scattered
or in small groups, covered by epidermis which later
crack, pale cinnamon-brown. Paraphyses cylindrical, curved
or bent, rarely slightly capitate, wall thin, sometimes
slightly thickened near apex. Urediniospores ellipsoid or
obovoid;,) 20-34) x 17-23) pm, wall. -1.5 pm thick,” hyaline;
echinulate. Telia hypophyllous, nearly black, crustlike.
Teliospores, in 3-4 layers, “ellipsoid, 23-30 x 14-16) um,
wall 1 pm thick, brown.
This rust, which is new to Kenya and Uganda, was
described on Vernonia sp. from Madagascar. The three hosts
are new to the rust.
Physopella nyasalandica Cumm., Bull. Torrey Bot. Club 87:
Say 19605
On Helichrysum gerberifolium Sch. Bip. (Asteraceae)
Masaka Distr., Kalungu Co., near Kasasa N of Mukoko, 1180
My Oo.UGet Sits hens puye Ch 108), £15
Uredinia hypophyllous, in small groups, surrounded by
curved paraphyses with a hyaline wall, slightly thickened
at apex. Urediniospores ellipsoid to obovoid, 21-27 x 18-
22 pm, wall pale yellowish, 1 pm thick, echinulate, pores
obscure.
The ‘rust is new to Uganda, previously known on H.
petersii Oliv. & Hiern from Malawi.
Physopella wiehei (Cumm.) Cumm. & Ramachar, Mycologia 50:
7a3y5 1958.
Syn. Angiopsora wiehei Cumm., Bull., Torrey Bot. Club 79:
ZS 9 be «
On Erlangea cordifolia (Benth.) S. Moore (Asteraceae)
Weomengo Distr.’, Kyadendo Co. ; Kazi; oi160%m, CG). 451), 22;
bisaco COiw,meike TOrests/ 111760) my (Gy (70) > ola.
On cf. Erlangea fusca S. Moore
EChiopias)) shoa Prov.,. (Wondo Gennet, 1800) my) 27. 111970;
Cy Ee EN
Uredosori hypophyllous, surrounded by more or less
incurved hyaline paraphyses, upper wall 2-3 pm thick, near
Bpex;) thickened”. to 8 (am lower wall .1-1.5 am thick.
Urediniospores ellipsoid or obovoid, rarely subgloboid,
19228) X° 16-22 pm, wall hyaline) 1=1.5 jam thick, densely
echinulate, pores not visible.
Apart from the type locality, Cummins (1952) pointed
out that the urediniospores described by Wakefield &
Hansford (1949) as belonging to Puccinia erlangeae Grove
on E. tomentosa from Kampala, belongs to Ph. wiehei.
Be cordifolia, jis a snéew host for this. rust
in Uganda, but it has previously been reported from Rwanda
(Majewski & Nowak 1982). E. fusca is a new host for the
rust which is new to the flora of Ethiopia.
PUCciInia BSCidisrormis Thuem., Flora’ 5820378, "1875.
On Microglossa pyrifolia (Lam.) O. Kuntze (Asteraceae)
Mubende Distr., Buwekula Co., Mile 124 on road Kampala-
Port Poncad, 2 1500. mG). o5):, LIL
Kenya. Eastern Prov., Nyeri Distr., Mt. Kenya, S slope,
Regati Forest Station, ©2200-2300 Mm, ~02.-03-.02°51973, “Ly
Ryvearcen AO }e, ) Opes Mire’ Non. Siqgon, 2100 me 27.80.1960;
Lind, III; Mt. Elgon, E. slope above Tweedie's sawmill,
ZoDuUN Ml, 22.02 01980," UO. Heaberg (85 7-UPS)). TaT.
Telia hypophyllous, in small groups, compact,
yellowish. Teliospores ellipsoid, constricted at septum,
58-74 x 22-33 pm, wall hyaline, smooth, 1-1.5 mpm thick, at
apex thickened to 4 pm. Pedicel hyaline, persistent.
518
Spores collapse when germinating.
This rust which is new to the flora of Uganda, is
previously known from Kenya, Tanzania and S. Africa, in
Tanzania on the same host. Other host genera are Nidorella
and ,senecio. lt lis simular tovP. Jalrata oH. .4) (eyes your
living on the same host, but the teliospores are somewhat
shorter, being up to 60 pm long and not thickened at the
apex.
Puccinia africana Cooke, Grevillea 8: 71, 1879.
On Spilanthes mauritiana (A. Rich.) DC. (Asteraceae)
W. Mengo Distr., Kyadondo Co., Makerere, 1200 mo,
03 UZ0196S Kae Lye (258), Gi. - (264) “land, Kabanvolo, tz00
m'Gy]. °° (492)3 Mawokota Co., ..Mpanga forest, 1150) m,AuGu
(178)3. Bugisu, Distr.) -C. -Bugisu. Co., near Busano, Vou0-
T1700 Vitti ok 2h. Ole tI6S. S KoA. bye. (166))) 2 aNeo Buc sutmconr
Bumasifwa, 1300 m,\ Gj. & P. Rwaburindore (407); Kigezi
Distr.) RubandasCov,— Kalinzu forest, 1400:.m, Giza (iso
all with Iilponly..
Kenya, ‘Central ‘Prov., " Mt. Elgon, — Suam Forest pStation,
2100 ,¢.234724. 010 | 923.0. Ryvarden (9259)5 721.
Telia hypophyllous in necrotic, brown leaf spots,
irregular, in groups, confluent or scattered, chestnut-
brown, greyish when germinated. Teliospores obclavate or
ellipsoid, constricted at septum, often falling apart, 37-
54 x 13-18 pm, wall 1.5 pm thick, thickened upward, at
apex up to 4.5 pm, yellow-brown in upper cell, going to
nearly hyaline at the base, smooth. Upper pore apical,
lower at septum, not visible in ungerminated spores.
Pedicel hyaline, persistent, short.
P. africana is widespread in tropical and southern
Africa, ‘and also, in Madagascar, (on, /S.4sacmellalwhurc
Wakefield & Hansford (1949) reported it on S. acmell'a from
several localities in Uganda. S. mauritiana is a new host
in Uganda, but it has been reported from Kenya and S.
Africa. Another host, in Tanzania, is Cineraria grandi-
flora Vatke.
Puccinia artemisiae-afrae sp.nov. (Fig. 4)
Telia hypophylla, plus minusve rotundata, in gregibus
irregulares vel in striis aggregata, pulvinata, brunnea,
post germinationem albo-grisea. Teliosporae clavatae vel
ellipsoideac,, sad. septa constrictdae, 55-90, x 316-27 eum,
pariete laevo, pallide brunneo, 1.5-3 pm crasso, apice aut
parum externus aut magis productus obtusissimusque, usque
14 pm crasso, poro superiore apicali, poro inferiore prope
septum, in sporas nondum germinatas invisibilis, pedicello
hyalino, usque 90 pm longo sed saepe breviore effracto.
Holotypet. Woe B.e) Gierum, 2375.) (MEU), ) 2310.1 970,9 sUganca,
Bugisu | Distre,)) Nea bugisu «Co.,. .-Sasa ~ Hut 73250" i eon
Artemisia afra Jacq. ex Willd. (Asteraceae). Isotype in
NPPI.
P. artemisiae-afrae which seems to be a_ leptosporic
species, resembles in many characters some of the forms
519
aggregated in the species P. cnici-oleracei Pers. ex
Desm., but the teliospore size eaSily separates it from
the latter species. Compared with P. artemisiae-sibiricae
Jerst., described from Mongolia, it is separated by its
larger and paler brown teliospores. The apex is not so
thickened and more regularly attenuated as in Ee
artemisiae-sibiricae.
Artemisia afra occurs in the more elevated areas from
ELniopia to South’ “Africa, ‘at’ “Mt, BLGensup stow 4100 om
altitude. From Natal, Transvaal and the Cape Province it
has been reported as a host for Pea tanhaceti) DC (svn.
absinthii (Hedw.f.) Dc.).
( JC ==
Erg. 24. Puccinia artemisiae-afrae he sp.
Teliospores from type.
520
Puccinia Dbakovarda Pate) S Har, ids .bOLe lhe Siu aiee eg eae
1900,
Synw sAccraium mitracarpi HH, | & PP. Syd... Annis mycol. 7h:
he nin he A
On Mitracarpus villosus (Sw.) DC. (Rubiaceae)
Teso Distr ., (Soroti'.Co., Arapai Research Station) 1100cm,
Gist \CATG ) ans
Aecia in groups, mainly hypophyllous, peridia white.
Aeciospores angularly subgloboid or ellipsoid, pale
yellowish, , 16-20 x 14-17 pm, wall 1 fim thick, verruculose
and with refracting granules.
Mayor (1964) who made biometrical studies of the
three rubiaceous rust species P. lateritia Berk. & Curt.,
PP. DOLESraae! Pj) 1& i. Syd. and P. bakoyana could not
separate them morphologically. Therefore, he placed them
all within P. lateritia, the first described one. Eboh
(1977) found that aecia on M. scaber, corresponding to A.
mitracarpi were displaced by telia matching those
described for P. bakoyana. He pointed out that the two
other species had longer spores, in P. borreriae also
strongly thickened at apex. As aecia are not mentioned for
the two other species, and the material studied by me
consists only of aecia, I prefer to follow Eboh and place
it within P. bakoyana.
Wakefield & UHanstord °(1949)) | reported, it wast
mitracarpi from Serere in Teso. The rust is widespread in
Tropical and South Africa on species of Borreria, Diodia,
Mitracarpus and Spermacoce. M. villosus is a new host for
the rust.
Puccipia -becii -Doidge, Bothalia/'2: 200)" 1926.
On Becium obovatum (E. Mey.) N.E. Br. (Lamiaceae)
E. Mengo Distr., Kyagwe Co., on the hills near Kimera, 1300
m, 11.09.1969, K.A. Lye, Rwaburindore & Faden (3844), I.
Aecia hypophyllous, in small, dense clusters. Peri-
dium revolute, white. Aeciospores angular, 14-18 x 12-16
pm, wall 1 pm thick, hyaline, delicately verruculose, and
with refracting granules.
This “rust),is,onewo tothe |) rust). flora “or jUganda:
described on the same host from Transvaal in S. Africa and
later reported from Malawi, and from there also on B.
odontopetalum C.H. Wright.
Puccinia cephalandrae Thuem., Flora, p. 425, 1876.
Syn. Aecidium cephalandrae Cooke, Grevillea 13: 6, 1884.
On Zehneria scabra (L.f.) Sond. (Cucurbitaceae)
Mubende Distr., Singo Co., Mile 101 on the Kampala-Hoima
FOad)) AOU mere i ON alsa ae le ese Ds,
Aecia hypophyllous, in small groups. Peridium white.
Aeciospores subgloboid or obovoid, 21-25(-32) x 17-23 pm.
Wall 1-1.5 mm thick, hyaline, verruculose, some spores
with refracting granules. Uredinia hypophyllous, round or
oblong, scattered or in small groups, cinnamon-brown.
Urediniospores ellipsoid or subgloboid, 23-28 x 18-25 ym,
BZ
wall 1 pm thick, cinnamon-brown, echinulate and with 2
equatorial pores. Telia in the uredinia, dark brown.
Teliospores ovoid or ellipsoid, with a pointed apiculus,
slightly constricted at septum, 37-47 x. 27-33 pm, wall 3-
4(-4.5) pm thick, brown, with ridges, best seen on dry
spores, upper pore apical, lower pore at septum. Pedicel
hyaline, deciduous.
The rust is new to the flora of Uganda, but it has
been recorded on several host genera in S. Africa, and it
is also known from Kenya, Zambia and Zimbabwe. Z. scabra
is a new host for this rust. nine
The Indian species P. cephalandrae-indicae H. & P.
Syd. differs from P. cephalandrae having urediniospores
with scattered pores and smaller, verrucose teliospores.
Puccinia crassocephali n.sp. (Fig. 5)
Pycnia, aecia et uredinia non visa. Telia hypophylla,
compacta, parva, rotundata vel irregularia, confluentia,
flavescentia. Teliosporae obclavatae, rotundatae vel
interdum ad apicem oblique attenuatae, ad septum leviter
constrictae, 33-50 x 10-17 pm, pariete, 1-1...5, pm crasso,. ad
apicem usque 6 pm incrassato, laevi, hyalino, poro
superiore apicali, poro inferiore obscuro (? prope
septum), pedicello hyalino, minus quam 10 pm longo.
Holotype: H.B. Gjerum 255 (MHU), 02.10.1970, Uganda,
Kigezi distr., Bufumbira Co., at the saddle between
Gahinga and Muhavura, 3100 m, on Crassocephalum aframonta-
hum R.E. Fr. (Asteraceae). Isotype in NPPI.
To my knowledge no Puccinia has been described for
the genus Crassocephalum. Puccinia species described for
allied host genera donot fit the P. crassocephali.
Nearest seems to be P. schizocarphae Jacks. & Holw.,
described from Guatemala and also reported from Colombia.
However, a comparison with the type (herb. kK) shows that
this species has more yellow sori and teliospores less
constricted at septum with a thicker apical wall than has
the new species.
40mm
Fig. 5. Puccinia crassocephali n.sp.
Teliospores from type.
522
Puccinia™desertorum H. & Pi. Syd.) “Bot. Vbe8 45 51 1259-=265-
1291075
On Evolvulus alsinoides (L.) L. (Convolvulaceae)
Kenya. Central Prov se, Thika Distr, 5 km WSW Or Thikeyewor
Hanis -Farm, wbolUUamsy 24401219705" Li) Ryvarden 5650." Ojm lie
CoastrProw.,) “lalcat Disth. ,) -lsavotNatrionaw ras, wast. sano
km ENE“Of Vol, .Ndolola, ..500 m, 08.0715 19707 WLS (Ryvarcen
(501-03 PO) Fadel.
Uredinia amphigenous, round to ellipsoid, sometimes
coalescent, brown. Urediniospores globoid to ellipsoid,
25-29) SX 222205 en, wall yellowish brown, distantly
echinulateé, =255=3 “um thick; “with two equatorial) pores
covered with low, hyaline papillae.
The rust, described on the same host from Namibia and
also reported from India, is new to the flora of Uganda.
Jerstad (1956) pointed out that P. desertorum "does not
appear to be very different" from P. lithospermi Ell. &
Kellerm., mainly reported from the Americas, but also from
the Philippines “and "India, “often found “on \the= “host
mentioned above.
A comparison of the type of P. desertorum with three
American specimens of P. lithospermi (all specimens herb.
S) showed that the teliospores of the latter were darker
brown than those of the former. Whether this difference is
Significant or not can probably be decided only from study
of more material from both continents, including also the
type of P. lithospermi.
Puccinia”’ dirformis Kuntze.in Kuntze %&\Schm., Myk. ~ Heflee,
TESS lege Onan
SynseP.- Vanbiguas (Alb. & Schw.) Lagh., ex*Bub./, Sitz. ber:
KY Bohm. Gess*’ Wiss 4,2 Maths nat. Cl... 1898s" 1477 1695.
On Galium spurium L. (syn. G. vaillantii DC.) (Rubiaceae)
Bugisu Distry, Ne oBugisu® Covent. Elogon7) banboos 1 Orecer
above Sasa River on? track to Sasa’ Hut, 3000 °m;, CG). 1(332)ea
sg A 2 lige
Aecia hypophyllous, cup-shaped and with revolute
peridia. Aeciospores angular, oblong or globoid, 20-25(-
30) x (13-)17-23 pm, wall 1 pm thick, hyaline, verruculose
and with refracting granules. Telia in streaks on stems,
on leaves hypophyllous, rounded. Teliospores cylindric to
obclavate, slightly constricted at septum, 42-60 x 18-22
pm, wall I-175, <am thick, . thickened at apex to jt) jim,
cinnamon-brown, brighter at apex and also in the _ lower
part of the spore. Pedicel nearly hyaline, persistent.
Telial sori and teliospores are not as dark brown as
they usually are, possibly due to juvenility.
This rust is widespread in Europe, Asia and North
America. In Africa it is known from Kenya and Algeria, and
also from the Canary Island and the Azores. The main host
is G. aparine. The rust is new to Uganda, but the host is
reported with the rust in Norway (Hylander et al. 1953, as
Gu vaillantit):
525
Puccinia dummeri Wakef. in Wakefield & Hansford, Proc.
Pent wSOC.. LONG. , CCSS. 100, Loao-s9 veto 2s 178, 1949.
On Echinops hoehnelii Schweinf. (Asteraceae)
Bugisu Distr., “N. Buogisu Co.;, near Sasa River on track to
SaseeHit., EZOUULNN WOT mts Oo) fonk t.
Uredinia hypophyllous, scattered, irregular, brown.
Urediniospores subgloboid, sometimes globoid or obovoid,
32-34. ee 51=37 pom, wall 1-1.5 pm thick, cinnamon-brown,
echinulate and with 3 equatorial pores.
This rust seems not to be known outside Uganda where
Lumbwa in Kyagwe is type locality. Type host is E.
aliplextcaulissOliv.. lt is also reported (on, EB. gigantel A.
Richy and Es sp.0b. hoehnelii*°is a new-host for the rust
species.
P. dummeri is closely related to P. pulvinata Rabenh.
and (Pe echinopis DC. (Wakefield & Hansford op. cit.). P.
pulvinata differs from P. dummeri in having urediniospores
with 2 slightly superequatorial pores. P. echinopis (=P.
calcitrapae DC.) has 3 equatorial pores. However, spore
eizes, £Or P. caleitrapac given in literature vary, but
they are mostly smaller than found in my material. Telio-
spore length is also shorter than given in the diagnosis
Lone. aummeri,
c
Puccinia erlangeae Grove, Bull. Misc. Inf. R. Bot. Gdns
KewiNo., 102) 2/0,.1916,.
On Erlangea cordifolia (Benth.) S. Moore (Asteraceae)
WeeMengowDistr.,. DUSIErO.CO.,..zikea, forest, liGO om, ~ 0G).
(WO DUG SUL DIStr. , N.) BUgisuCo., on track to Sasa
Hut, below Bulambuli Escarpment, 2000 m, Gj. (409), III.
Ong bee Ces LUSCa
SUGISULDAstrey N. BUgGISU CO, ,ptoasa Hut, 3250 mM, Gy. (372),
aba
Uredinia hypophyllous, brown. Urediniospores subglo-
Doig, .ellipsoid or obovoid, § 24-32 x 21-24 ym, wall 2-225
DieechscK,,, densely echinulate aniduwith 2. distinct, sequa-
torial pores. Telia amphigenous, mostly hypophyllous, oval
or oblong on veins, brown, pulverulent. Teliospores 31-41
x 24-30 pm, broadly ellipsoid, mostly rounded at both
ends, slightly constricted at septum, wall (1-)1.5-2(-2.5)
pm, pale brown, upper pore apical, lower near the hyaline
pedicel.
The teliospores on E. cordifolia are slightly shorter
than.are those on E. fusca and with a thicker, wall,- but
the form, colour and echinulation are similar.
Wakefield & Hansford (1949) have previously reported
this rust on E. tomentosa S. Moore from Bufundi in Kigezi
and from Kampala. The urediniospores described from the
specimen collected at Kampala, Cummins (1952) pointed out
as belonging to Angiopsora wiehei Cumm. (= Physopella
wiehei (Cumm.) Cumm. & Ramachar). The description of the
urediniospores given by Henderson (1970) from Tanzania
Matches well with the urediniospores in the present
material.
EB. Cordiftolieawis a new host record for” this rust.
524
PucciniaquizotiaeCumm. , Bull. Tormey Bot sClub 7 os ii2 21
1oS2 6
On Guizotia scabra (Vis.) Chiov. (Asteraceae)
Teso Distr., Serere Co., Serere Research Station, 1150 m,
She MASS) i pees BunyOro.. DA str. .Budahya Cow UnOLmMear |) loo
Tipe) Gael SS Wy aioli.
Telia hypophyllous, in dense clusters, compact,
cinnamon-brown. Teliospores oblong-ellipsoid or obclavate,
constricted at septum, 41-57 x 13-21 pm, | wall 1-1:.5) jim
thick, at apex thickened to 6 pm, yellowish to pale brown,
smooth, upper pore apical, the lower near septum, obscure
in ungerminated spores. Pedicel hyaline to pale yellowish.
One-celled spores occur.
The rust was described on G. scabra from Semuto Road
in E. Mengo. Later it was reported on the same host from
Lake Mulehe in Kigezi by Ebbels (1972), and on which it is
also known from Kenya and Sudan. From Ethiopia it is
reported on .G. “abyssinica’ (set. ) Cass.) “and GliwischuLecas
Hochst.
Pucciniaihansfordiana iCumm... Bul. Torrey (Bot. (Clap iy ue
PARE MTs We YAR i
On Senecio trichopterygius Muschl. (Asteraceae)
Kigezi Distr., Rubanda Co., Mile 26 on the Kabale-Kisoro
roads) welOOam, wiGhe 207)» jLoraRuobanaGa Cos, ECRUVa. fore ams
Mile 34-35 on the Kabale-Kisoro road, 2100 m, Gj. (228),
Tf, Bufumbira’Co.}") on. |,track to Gahinga,, below) Fores:
Reserve Border, 2500 m, and at the saddle between Muhavura
and Gahingea ci ot OO Mp Gt eh Ome oi, 2 OF en.
Uredinia mainly hypophyllous, round or oval in face
view, scattered or in groups, brown. Urediniospores ellip-
soid,)|) obovoidor./subgloboid, 27-37 .x\:23-30 pm wall teo-2
pm thick, dark cinnemon-brown, echinulate, with 3 mainly
equatorial pores.
This) rust species was described on S. denticulatus
Engl. from Kachwekano Farm in Kigezi. Wakefield & Hansford
(1949) reported the same specimen.
S. trichopterygius is a new host for the rust.
Puccinia” hedbergiaeL,) Holm, |) Svensk bot! Tidaskr. 76702 oo,
AOTS'.
On Senecio erici-rosenii R. & Th. Fries (Asteraceae)
Kigezi Distr., Bufumbira Co., Mt. Muhavura, W slopes, 3400
mM, US SO M1940 7 Of Hedberg: (Z05660 UPS }yeLlll.
Telia amphigenous, irregular, scattered, pulverulent,
Cinnamon-brown. Teliospores ellipsoid or obclavate,
slightly attenuate at apex, constricted at septum, (33-)
40-57 x (15-)17-23 pm, wall 1-2 pm, at apex thickened to
3 jam, pale cinnamon-brown, smooth, pores obscure. Pedicel
hyaline, long, persistent.
This rust was described on Senecio meruensis Cotton &
Blakelock from Mt. Meru in Tanzania. It is new to the rust
flora of Uganda and S. erici-rosenii is a new host for the
rust. Both hosts belong to Dendrosenecio. In the Uganda
525
specimen some spores are longer than in the diagnosis (38-
48 x 20-25 pm).
Puccinia holosericea Cooke, Grevillea 10: 126, 1882.
On Ipomoea hederifolia L. (Convolvulaceae)
Way Mengo Distr .),))).Busiro Co... Entebbe, near the Botanic
Garden, WlSs0hm Gayo) (CLS) yi Lil; Bunyono Distr .;\\Bujenye
€o;.,, Nyabisabu mear Nyabyeya, 1100 m, Gj). (352),)TIL.
On Ipomoea involucrata Beauv.
Kigez) Distr.) Rubanda)Co., \Nyamabala 2100) m, Gj. (226),
ELI.
On Ipomoea wightii (Wall.) Choysy
Ankole: Distr), BunyaruguruCo., Kalinzu: forest) 1400 m,
Grint vay yt BIT.
On Ipomoea sp.
Kenvyawm Eastern Prov., ‘Nyeri) Distr.) Mt. Kenya, |S): slope;
RhRegqammerOrestrotation, | 2200 —. 2300 m, o2.-352.1973, Ls
Ryvaroen, (91094) Oj) TTT.
Telia hypophyllous in necrotic spots, arranged in a
circle around a central sorus, compact, cinnamon-brown to
chestnut-brown. Teliospores oblong, ellipsoid or obcla-
vate, slightly constricted at septum, rounded at apex,
rarely attenuate or truncate, 33-54 x 15-20(-23) pm, wall
Wi5-29m thick, ‘vat apex jthickened, tos pm,)) smooth, | pale
Cinnamon-brown. Pedicel hyaline to pale cinnamon-brown,
persistent. One-celled spores occur.
Wakefield & Hansford (1949) reported this rust on
Convolvulaceae (in the register is said Ipomoea) from
several places in Uganda. In Africa it is reported from
Several 1.) spp. including. I. wightii.,1.. hederifolia and
I. involucrata are new hosts for the rust which also
occurs on Lepistemon owariense Hall.
Puccinia hoslundiae H. & P. Syd. emend. Vien.-Bourg.,
Uredineana 4: 175, 1953.
Syl Pe ihoslundiacuh. f'P. Syd..in: peutsch. Zentr...Atxika
Bexped. 1907-8 fi22enr97,, 19149.Po\hoslundiae Grove; Bull.
esc went. Rie Bot. (Gdns, KewiNo. 100 270) obs.
On Hoslundia opposita Vahl (Lamiaceae)
Teso Distr., Serere Co., Serere Research Station 1150 m,
GHeCs SOL TO) IE +077
Uredinia amphigenous, rounded, scattered)! ori) an
groups. Urediniospores obovoid to subgloboid, 24-28 x 18-
24 pm, wall yellowish brown (1.5-)2-2.5 pm thick, echinu-
late, with 2 equatorial pores, often obscure. Telia hypo-
phyllous, rounded, dark brown. Teliospores ellipsoid, 28-
43x 22-24 pm,)iwall 3-3.5 pm thick, -at apex thickened to 6
um, finely verrucose. Pedicel hyaline, collapsing, in some
spores as long as the spore, but mostly broken shorter.
Wakefield & Hansford (1949) who described the aecial
stage of this rust, reported it from W. Mengo and Kigezi
Districts. .It is known on the same host in Sudan, Ghana
and) ivory | Coast. | Type host\is)H. verticillata Vahl,
described from Rwanda.
526
Puccinia isoglossae Doidge, Bothalia 2: 202, 1926.
On. HypoGstes verticillaris (L.f.) Soland ex" ‘Roem. &
Schult. (Acanthaceae)
Kabamoj}arDistre;, PMi.! YMoroto; 2/700=sl00rmp 26.2210 600 7 me
Long -Brauny (60 j) 2rb.
Telia iypophyllous, rarely epiphyllous}. (compact, in
very dense groups, dark cinnamon-brown. Teliospores
fusiform or cylindrical, slightly ‘constricted ‘at septum,
32-42 x 10-13(-16) pm, wall 1 pm thick, at apex thickened
to 4 pm, hyaline, smooth. Pedicel hyaline, persistent.
P. isoglossae has been found scattered over Africa
from’ (‘Kenya fand@Stdan) to S ;)OAfrica: On “Hal iverticiiianmas
(originally described on Isoglossa ovata Nees and I. woodi
C.B.Cl., but hosts wrongly identified — “cf. Doidge 194538
It was described on H. cunninghamiana Benth. & Hook. from
Taiwan as P. hypo€stis Sawada in 1943, but not validly
published (ct. Waundoni'963')*.
Pucciniakaichbrenneri: de Toni, “Sylils (Fund./2) 66579-1860"
var. kalchbrenneri.
On Helichrysum globosum Sch. Bip. (Asteraceae)
Ankole Distr.),,"°Bunyaruguru Co.) Kalinzw Forest,*’ nean che
Ssawnickiy \TA00-mi Gil. (144), II; Kigezi Distr., Rubanda
Co.) Lakes Bunyony i Sharp's ‘Tstand /-71:800 myiiG).." C2338)
Rubanda Co., Mile 28 and Mile 34-35 on the Kabale-Kisoro
road, §*2700°m,.., (G3) (202 “and 229) fll, Kabale, 1o00™m pea.
(21:7) polis IDL Kinks 2a PCoe, Bwindi, Gyt (233) 9°11 elie
Uredinia hypophyllous on lower side of yellow-brown
spots, scattered or in small groups. Urediniospores
obovoid to subglobose, 24-30 x 19-22 pam, wall 1 pm thick,
pale yellowish, densely echinulate, pores obscure. Telia
hypophyllous, dark brown. Teliospores ellipsoid or
clavate; constricted) at: septum, »/53-62 x 23-27 Gum, Powel
yellow-brown, at apex nearly hyaline, 2 pm thick, at apex
thickened up to 10 um, pedicel short, deciduous.
Wakefield & Hansford (1949) reported this on H.
globosum,) Hainudtetorum: DC.) He undatum “Less. jiand of. ‘spp.
from many localities between Mt. Elgon and Kigezi. This
rust is also reported from Ethiopia, Malawi, S.. Africa and
India.
The var. valida Doidge, described from Transvaal has
longer spores, up to 80 um, and with apex thickened up to
3. namie
Puccinia lantanae W.G. Farlow, Proc. Amer. Acad. Arts Sci.
184, Bare RO SS hee Lon 6:)
On Barleria sp. near grantii Oliv. (Acanthaceae)
Kenya. Eastern Prov., Nyeri Distr., Mt. Kenia, S slope,
Regati “Forest (Station," 72200-2300) im, 702 03 7022197 S,aeeee
Ryvarden (9119, O).
Telia hypophyllous, small, round, often in dense
groups, compact, brown. Teliospores light brown, two-
celled spores ellipsoid or clavate, slightly constricted
at septum, mostly rounded at the ends, but sometimes
AN |
attenuate at apex or tapering to the pedicel, 27-40 x 15-
Zio ite) WallesiiS um Chick, sehickened,to..8 um at, apex,
smooth. One-celled spores common, ellipsoid, light brown,
£7 20g x No- 2) um. pedicel hyaline.
P. dlantanae was originally described on species of
Lantana from the Americas. Laundon (1963) found that
several other rusts described on hosts belonging to
Acanthaceae showed the same feature and made them = synony-
mous with P. lantanae, and indicated that P. heterospora
Moncsalso, be dncluded. \ InVAfirica the rust, jis’ reported jon
Lantana amara from) Ivory Coast and Ghana, -and’ on L. sp.
from Ethiopia. Barleria is a new host genus for this rust.
Another African rust on this genus is Aecidium barleriae
Doidge.
Fig. 6. Puccinia lantanae
Teliospores (from Ryvarden 9119, O)
528
PucciniailaterdtialRerks 8 Curton wo. Aegean Nat. sce
Philad.))\ New iSert 22281, {VE53%
On Spermacoce ocymoides Burm. f. (syn. Borreria ocymoides
(Burm. f£.) DC.) (Rubiaceae)
Bunyoro Distr., Bujenje Co., Budongo Forest, 1050 m, Gj.
C3 5Q0, RB.
On Spermacoce princeae (K. Schum.) Verdc. (syn. Borreria
princeae K. Schum.)
W. Mengo Distr., Mawokota Co., Mpanga Forest, 1150 m, Gj.
CUTS bie) Bunvore Distr. i BugahvarCow.| HoimaysmoUGeim.
ey BU, (i WY40 Op Eee Us Ba
On Spermacoce pusilla Wall. (syn. Borreria pusilla (Wall.)
DG)
E. Mengo Distr., Buruli Co., Mile 125 from Kampala on the
Masindi» road; 91250) m, Gz. (356), .22R, . Mubende’ Distre;
Singo Co., Mile 112 from Kampala on the Hoima road, 1000
me) Gist 22a yh LEE
On Spermacoce ruelliae DC. (syn. Borreria ruelliae (DC.)
H. Thoms)
E. Mengo Distr., Bulemezi Co., Nandere, 1200 m, 24.6.1960,
Katete, III.
Telia hypophyllous on brown, nectrotic leaf spots.
Sori rounded to oval, often confluent, compact, brown,
becomming whitish when the spores germinate. Teliospores
wide ellipsoid or obovoid, not constricted at septum, 27-
37 x 15-24 um, wall 2.5-3 pm thick, at apex thickened up
to 7 um, smooth, hyaline to pale cinnamon-brown, rarely
Cinnamon-brown. Upper pore apical, the lower near septum.
Pedicel persistent, hyaline, sometimes obliqually attached
to the spore. One-celled spores occur.
Wakefield & Hansford (1949) reported P. lateritiae on
S. ruelliae (as B. ruelliae), S. senensis (Klotzsch) Hiern
(as; B. sinensis: K.Schum.) jand) on) Borreria» sp.) 7prom
several localities in Uganda. It is known from many
countries..in, Africa, and also from India. Other | hest
genera are Dioidia and Octodon. S. princeae seems to be a
new host for the rust.
Puccinia leonotidicola P. Henn. in Baum, Bot. Ergebn.
Kunene-Zambesi Exp. p. 157, 1903.
On Leonotis nepetifolia (L.) Ait.f. (Lamiaceae)
W. Mengo Distr., Kyadondo Co., Kampala, Makerere Hill,
1.200. m,, 280830970, Hak Tt Tatigoola, ill, Gi. (62606) ae,
Kabanyolo, 1200 m, Gj. (281), II, Mawokota Co., Nakysanja
(Kawanda), 1180 m, 08.11.1970, J. Mukiibi, II, Teso Distr,
Serere Co., Serere Research Station, 1150 m, Gj. (429), II.
Kenya. Eastern Prov., Nyeri Distr., Mt. Kenya, S. slope,
Regati Forest Station, 2200-2300 m, 02.-03.02.1973, UL.
Ryvarden (9114, O), II.
On Leonotis sp.
Tanzania. Moshi Distr., Mt. Kilimanjaro Reg., between
Lemosho and Shira Plateau, 2900 m, 21.01.1970, K.A. Lye,
ee
Uredinia amphigenous, irregular, scattered, often
covering the lower side, brown. Urediniospores subgloboid
529
to globoid, rarely ellipsoid or obovoid, 23-30 x 22-28 pm,
wall 1 pm thick, cinnamon-brown, echinulate and with (3-)
4-5 scattered pores.
The rust is widespread in warmer areas in the
Americas from Paraguay to Florida, in Africa, and also in
India. In Uganda Wakefield & Hansford (1949) reported it
OnyL. (mollissima) Gunge, L. | nepetifolia:and L. ‘sp. from
several localities.
Puccinia Leucadiss Pi a He isyd.) . Monogr: Uredi’ 'he"281,
1902:
On Leucas calystachys Oliv. (Lamiaceae)
Mubende Distr., Singo Co., 10 km NW of Katera, 1400 mo,
16.03.1969, K.A. Lye (2306), Lester & Morrison, II.
On Leucas martinicensis (Jacq.) Ait.f.
W. Mengo Distr., Kyadondo Co., Kampala, Makerere Hill,
E200 SO 10.1968, K.AL Lyé (12), ‘Il, Kabanyolo;, 1200 m,
Gj. (488), II, Mawokota Co., Mpanga Forest, 1150 m, Gj.
ChSO),. Els) Mubende Distr. , Singo Co.) Kiboga, “1300°m, Gj.
tour), tl.) BUNYOrO, Distr. , Bugahya Co.) Holma, 1500 °m,. G4.
os0) simile ih) wage MDISstr se . | BuLumbiray Co, con: track. J.to
Gahinga, just below the forest boundary, 2500 m, Gj.
(244a), II.
Uredinia amphigenous, rounded or oval, cinnamon-
brown. Urediniospores mostly globoid, 25-31 x 24-31 pm,
wall 2-3 pm thick, cinnamon-brown, distantly echinulate
and with 3-5 pores.
The rust is widespread in Africa, and it is also
reported from the Dominican Republic, India and China.
Wakefield & Hansford (1949) reported it on both hosts
mentioned above from a few localities from the Mt. Elgon
area, to Kigezi«
PrecantarruppLicola Pate, & Harry; Bulls Soc. Mycol. Fr: 24:
roy. Loo,
syn. P. tlippiivora H. & P. Syd. in’Wildem. Flore Bas- et
Mover Congo sj rasc.. T2117 19093) P. natalensis Diet. & "HH.
Syd. var evansii Doidge, Bothalia 2: 82, 1926.
On Lippia grandiflora A. Rich. (Verbenaceae)
MerhNengo Distr., ‘Bbusiro Cco., 3) km-SE-“of Jungo Church, 01.40
mene s “ove Cu9t2Z) & Mo) Morrison, Itl;* Bunyoro) Distr,
Bunyorg) ‘Ranching Scheme Area, 91100 mp 327.09.196T, "Bos
turner: (209); Lil.
On Lippia plicata Bak.
Tanzania. MIringa Distr., Ruaha National Park, Magangwe
Ranger Post, 1330 m, 09.03.1972, leg. A. Bijgrnstad (1461,
34 fain 8 i Be
Telia hypophyllous, Sound cor oval, pulvinate,
cinnamon-brown, greyish when germinated. Teliospores
ellipsoid, not constricted at septum, (34-)43-52 x (16-)
19-26 pm, wall smooth, 1.5-3 pm thick, often unevenly, at
apex thickened to 5.5 pm, pale yellow, upper pore apical,
lower near septum, obscure. Pedicel hyaline, persistent,
longer than the spore. One-celled spores fairly common.
P. Jlippiicola has been reported by Wakefield &
Hansford (1949) on L. adoensis Hochst. from Kipayo in
Kyagwe and Serere in Teso and on L. sp. from the Elgon
area. Outside Uganda it is reported on Lippia spp. from
Ghana, Zaire and Kenya southwards to South Africa. _L.
plicata appears to be a new host for this rust species.
Puccinia menthae Pers., Syn. Meth. Fung. p. 227,1801.
On Satureja pseudosimensis Brenan (Lamiaceae)
Kigezi Distr., Rubanda Co., Nyamabale, 2100 m, and Echuya
Forest: Reserve, ) Mile 33,, 2100 m, 2G}. /°( 219) -239a) alee
Bufumbira Co., Forest Reserve on track to Gahinga, 2700 m,
Gi. 248), °LIx
On Satureja punctata (Benth.) Briq.
Kigezi Distr., Rubanda Co., Echuya Forest Reserve, Mile 33
Exot Kalsay2i0eme Gaca( 239), 1 2
On Satureja simensis (Benth.) Briq.
Kigezi' Distr.,..Rubanda Co.,. Mile 26. from-Kabale on ‘the
Kabale-Kisoro, road; 92100 m,..Gy}. (206), 10
Uredinia hypophyllous, round, oblong, confluent.
Urediniospores ~obovoid, (or ellipsoid, °§22-26 xv10-Zimam,
wall 1-1.5 pm thick, pale cinnamon-brown to hyaline, echi-
nulate with 2-3 equatorial, often obscure pores.
The only record of the worldwide distributed mint
rust, living on a number of host generea of the Lamiaceae,
was given by Wakefield & Hansford (1949) reporting it on
Mentha aquatica L. from Kampala. The hosts mentioned seem
to be new for the rust.
Puccinia natalensis, Diet....H.. & PP. Syd.,,~Hedwigia® goa,
Beihert S730, 0 699,
On Lantana trifolia L. (Verbenaceae)
EY Mengo Distr<y “Bulemezai “Co.., | ‘Kiziba, §°1200'm; > Mugewys
(34); “Bunyoro: Distr: , ‘Bugahya: Co<, ‘Hoima, 111500 im, Ga
C325) Ankole Distr., Nyabushozi Co., Muko Research
Station / 1370) mow Gi O19), eal Lewithe Litaonly.
Kenya. E. Prov., Nyeri Distr., ‘Mt. Kenya, S. slope, Regati
Forest Station, 2200-2300, 02.-03.02.1973, Le. Ryvarden
69'1 28); sO} 2
Telia hypophyllous, rounded, pulvinate, dark brown.
Teliospores ellipsoid to obovoid, slightly constricted at
septum, , jrarely W cylindric, 50-68) x, (22-) 30-34) fun awa
yellowish brown, smooth, 2.5-3.5(-4) pm thick, at apex
thickened to 9.5 pm, upper pore apical, lower pore near
septum. Pedicel wide, up to 170 pm long, hyaline.
This rust, described from S. Africa and later repor-
ted from Sudan and doubtfully from Ghana by Hughes (1952),
has been recorded from several places in Uganda by
Wakefield & Hansford (1949), also on L. trifolia. A record
by Petrak (1959) on Lantana sp. from Tanzania is based on
a misidentificationof.the host (cir. Gillisj97zZ).
Puccinia necopina Grove, Bull. Misc. Inf. R. Bot. Gdns Kew
So
Noe 110217269-272), 1.976.
On Tristemma incompletum R. Br. (Melastomataceae)
W. Mengo Distr., Mawokota Co., 2 km E of Bujoho on Mubende
Road, sihsO omy KEA. Give (1946), Lit.
Telia hypophyllous, densely aggregate, yellowish
brown. Teliospores ellipsoid or clavate, constricted at
Ssepeum, 4134) 38470 xe GTO) 1 4—te nm; es wall ot” om tthick,
hyaline or very pale yellowish, smooth, pores not visible
in ungerminated pores, upper pore apical, lower probably
near septum. Pedicel hyaline, short.
P. necopina was described from Kipayo, and was later
reported by Wakefield & Hansford (1949) from Kampala, in
both cases on Tristemma sp.
Puccinaa ocimi ‘Doidge, Botha lila i2: 203) 1926.
OV eee oDum OCimi Ps Henn., Bots Jb. %/: Ve, 1893.
On Ocimum suave Willd. (Lamiaceae)
Kenya. Eastern Prov., Nyeri Distr., Mt. Kenya S slope,
Regati Forest Station, 2200-25002 Mye {2c oe tlio ioe tlie
Ryvarden ((91237° 0}, Eid.
telia hypophyllous, irregular, confluent, © compact,
dark brown. Teliospores clavate to ellipsoide, slightly
constricted at septum, 42-55 x 15-21 pm, wall (2.0-)2.5-
wei Chick, Imateapex thickened’ to 85 jm; »golaen brown,
smooth, Pedicel hyaline to very pale yellow, long,
persistent.
P. ocimi was reported on O. suave by Wakefield &
Hansford (1949) from Kyasoweri in Elgon and from Serere in
Teso, on O. sp. from Kampala while Ebbels (1972) reported
it from Lake Mulehe in Kigezi. It is widespread on Ocimum
Spee naar Cay = trom Ethiopia and Sudanuto: S. Africa.
Outside Africa it is reported from Madagascar, Comoro
Islands and India.
Puccinia pentadis-carneae Wakef. in Wakefield & Grove,
Kew Bull. 74, 1916.
On Pentas lanceolata (Forsk.) Deflers (Rubiaceae)
Kigezi1) Distr.,,"Rubanda Co., Nyamabale, 2100 m, G3... (220),
fib Bi
téelzva whypophyllous, round, “scattered ‘oriin, wdense
groups, compact, cinnamon-brown, when germinating paler.
Teliospores fusiform, sometimes eylLindric, slightly
constricted at septum, 33-44 x 12-14 pm, wall 3-4 pm
thick, at apex thickened to 8.5 pm, hyaline, smooth, upper
pore apical, lower pore near septum. Pedicel hyaline,
persistent.
This rust was described on Pentas carnea Benth. from
Kipayo in Kyagwe and later reported from Kinyala in
Bunyoro on the same host. P. lanceolata is a new host for
thrserust.
RUC eearmubtvcolalPavewH, Syd.) Monogr. Ured. hs) 226;. 1902.
On Rubia cordifolia L. (Rubiaceae)
a2
Kenya. Centrad Prov. ,|\\Thika Distr., ‘5S ‘km WSW#oL Thika,vac
B. Harris Fann, stS00um,y 20. 1970, DA cRyvardenaloozoy OT
CRRD Svike (yl AS Ei
Uredinia amphigenous, long covered by epidermis,
irregular, cinnamon-brown. Urediniospores subgloboid to
broadly ellipsoid or obovoid, 28-37 x 22-32 pm, wall 2.5-3
pm thick, yellowish brown, echinulate and with 2 (rarely
3) equatorial pores covered by low, hyaline papillae.
Telia epiphyllous, in uredinia, dark brown. Teliospores
ellipsoid or clavate, rounded or attenuated at apex,
slightly constricted et septum, (43-4) 54-62) 219-26) Bom,
Wall \\W. 542.) amy (thick)! at apex thickened) up) wo) Sig,
yellowish brown, but nearly hyaline at apex, smooth, pores
obscure. Pedicel jup to 75 pm long, hyaline,
P. xrubiicola was described from Ethiopia and later
reported by Wakefield & Hansford (1949) from Bufundi in
Kigezi on the same host. It is new to Kenya. Mogk &
Hindorf (1971) have reported Puccinia collettiana Barcl.
on P. cordifolia from Kenya. This species which) is repor=
ted from many localities in Asia from Pakistan, India,
Nepal and China, and also Kirgezia and Kazakhstan in USSR,
has a thinner urediniospore wall, by Nevodovsky (1956)
measured to be 1-1.5 pm thick. This corresponds with what
I have seen in specimens from Pakistan and India. It has
also darker brown teliospores. P. dimorpha P. & H. Syd.,
described from S. Africa but also reported from Uganda on
R. condifolia, has thinner urediniospore wall (1.5-2 pm,
cfr. Doidge 1926) with 3 pores and longer teliospores.
Pili rublivora PB. Magn.,); iknow only on sR. |) Trutvoose
Ait. from the Canary Islands, has teliospores not or only
slightly thickend at apex.
Puccinia) tuelliae 1H, “& P. Syd.,; Monogr. Uredi) Te) 2367902
Syn. P. ruelliae (Berk. & Br.) Lagh., Troms@ Mus. Aarsk.
17: 71, 1895, (III not described): Uredo ruelliae Berk. &
Boi, Us Lanne SOC. nbOt.) LOR A Siti OZ ators.
On Ruellia praetermissa Lindau (syn. R. patula Jacq.)
(Acanthaceae)
Bunyoro, Distri;) | Buyenje.' Co. Bukumi, 750 m,. Gj." 43 45)gees
alt a
On Ruellia sp.
Bunyoro'|Distr.,. Bujenje Co.);) Bukumi;)) 750 m,) Gyo (S48 jy 1ae
Uredinia amphigenous, scattered or in rings around a
central sorus, cinnamon-brown. Urediniospores globoid to
subgloboid, 23-33 x 21-28 pm, wall 2.5 pm thick, cinnamon-
brown, echinulate and with 2 equatorial pores. Telia
amphigenous, irregular, pulverulent, blackish brown.
Teliospores ellipsoid, not constricted at septum, 37-48 x
30-33 pm, wall 2.5-4 pm thick, chestnut-brown, verrucose.
Pedicel hyaline, often obliqually attached to the spore.
In the paper on rusts on Acanthaceae, Laundon (1963)
listed several Ruellia spp. as hosts for the present
species, and also Asystasia spp. and Eranthemum variable
Rw! ‘Br. in Africa vandijiAsia,’ In Africa! it, is’ Preportedcs irom
Ethiopia,’ ‘Ghana )iand); the’ ivory Coast)’ but/iti2s\ inews gra
559
Uganda.
Viennot-Bourgin (1953) following Arthur (1934) dis-
cussed this rust under the name P. ruelliae (Berk. & Br.)
Lagh. including some American species which Laundon
(op.cit.) maintained as separate species. A _ uredinial
stage on Lepidagathis laguroidea T. Anders from the Ivory
Coast, by Viennot-Bourgin (1958) placed within the same
species, Laundon (op.cit.) listed as Uredo sp. lLaundon
also discussed the nomenclature.
Puccinia semiloculata Laundon, Mycol. Pap. 69: 61, 1963.
On Justicia sp. (Acanthaceae)
Kenya. Eastern Prov., Nyeri Distr., Mt. Kenya, S slope,
Regati Forest Station, ZA00 52300) OMG cen slale, wi Lot Oipet, sae
RYVVarOenet Ot 247 Opie I re rT T
Aecia hypophyllous, in small groups. Peridia white,
revolute. Aeciospores ovoid, subgloboid or angular, 16-22
x 12-18 pm, wall 1 pm thick, hyaline, verruculose and with
refracting granules. Telia hypophyllous, small, rounded,
confluent, brown. Teliospores cylindrical, clavate or
fusiform, slightly constricted at septum, attenuate, rare-
PyeecuucaLe at apex, 36-49 x-i2-16 pm, wall nearly hya-
lane, ..smooth, 1l;pm thick, at apex thickened up to 7 pm.
Pedicel hyaline, persistent.
The following specimens have only aecia. According to
Lavndon ()963) PP. semiloculata and P. -thunbergiae” “M.C.
Cooke cannot be distinguished when lacking telia. The lack
of pycnia separate them from Aecidium justicia P. Henn.
On Justicia flava (Forsk.) Vahl.
WeeeMengo Dastr.s,, \Busiro Co., .Entebbe,. (1150 .m,\ Gi}. (17),
PyadOnUOn OG.) Kazi, tt o0 lm,» Gy. (55) 2) Bugisu Distr., Ne
Bagisu,Co., Bulambuli Escarpment, 1700 m, G}., (396).
Kenya. Central Prov., Mt. Elgon, Suam Forest Station, 2100
Mees s—-24.0e1 9/3, Le Ryvarden, (9275, 0).
On Justicia exigua S. Moore
Teso Distr., Soroti Co., Arapai Research Station, 1100 m,
Gi, 4419).
On Justicia striata (Klotzsch) Bullock or J. unyorensis S.
Moore
W.- Mengo Distr., Mawokota Co, Mpanga Forest, 1150 m, Gj.
(ies 1b.) Menge. Distr, Burulda Co. , Sof Katugo,1150%m,
Sieur ise);
On Justicia stricta (Klotsch) Bullock
W. Mengo Distr., Busiro Co., Entebbe 1150 m, Gj. (30), and
Zikalrovessu, 1 160.m, Gyiei (72):
On Justicia sp.
Bugisu Distr., N. Bugisu Co., Forest Reserve Boundary on
track, to’ Sasa Hut, 2500. m;, Gj.) (394).
Puccinia stobaeae Mac Owan var. woodii P. & H. Syd.,
Monogr. Ured. 13, 7160, 1902;
On Berkheya spekeana Oliv. (Asteraceae)
Karamoja Distr., Matheniko Co., SE of Sogolomon, Mt.
Moroto, 2300 m, 11.06.1970, K.A. Lye (M 18A), O+1I; Mubende
534
Distr..,,) SingovCo..,- Mile. 70° on. the Hoima road ya) 4000n,. Ga.
(298). ILstPiz.Bunyoro Distr. ,).Bugahya Co.,@Hoima, who00m.
Gy i UA0S23 PPR sere Lite toro Distre j KyakaCos, aiimer i oman
ForG Portalroadyo00) m,7Gi.% (49S y ibe
Pycnia epiphyllous. Aecia hypophyllous, in small
groups. Peridium white, long, revolute, lacerate. Aecio-
spores subgloboid, ellipsoid or irregular, 24-38 x 20-31
jam, wall 1 pm thick, hyaline, with large prismatic warts.
Uredinia mainly hypophyllous, cinnamon-brown. Uredinio-
spores subgloboid, 36-42 x (30-)33-38 pm, wall 2.5-3 jam
thick, yellowish brown, with scattered, long spines, and
3-5 pores with hyaline papillae, equatorial or scattered.
Telia hypophyllous rounded or oval on veins, pulvinate,
dark brown. Teliospores ellipsoid with rounded ends, rare-
ly obovoid, slightly constricted at septum, 40-52(-64) x
32-40 pam, wall cinnamon-brown, in lower cell 1.5-2 jam
thick) (vin upper cell 25-3 im, Yat “apex thickeneds tomean.
Am, finely verrucose, rarely smooth, upper pore apical.
Pedicel long as the spores, but often broken short under
the spore.
This: rust is’ “new to Uganda. ‘Both varieties rare
previously known only from S. Africa. The var. woodii has
urediniospores with thinner wall than has the Var.
stobaeae. The teliospores of var. woodii are shorter than
those of var. stobaeae, and with a wall distinctly thicker
in the upper cell than in the lower one while they in the
var. stobaeae are of evenly thickness.
Fig. 7. Puccinia tetradeniae n.sp.
Teliospores from type.
530
Puccinia tetradeniae n.sp. (Fig. 7)
Pycnia, aecia et uredinia non visa. Telia hypophylla,
parva, castaneo-brunnea. Teliosporae obclavatae, ellipsoi-
deae vel irregulares, ad septum leviter vel non constric-
tae, 43-60 x 23-35 fam, pariete 1-1.5 jim crasso, ad apicem
usque 8 pm incrassato, brunneo, laeve, poro superiore
apicali, poro inferiore prope septum instructis, pedicello
hyalino, usque 80 pm longo, hyalino, deciduo.
HOLOLY pe: ~S.BasuManum 7 1-12.(MHU) >). 12.07<61971,. Tanzania;
Distr. Njombe T., Nyantyla River, on Tetradenia riparia
(Hochst.) Codd (Lamiaceae). Isutype in NPPI.
To my knowledge no rust has been described neither on
this genus nor on related genera in the Stachyoideae -
Podostemoneae tribe (Engler & Prantl System).
Puccinia thunbergiae M.C. Cooke, Grevillea 10: 125, 1882.
On Asystasia gangetica (L.) T. Anders. (Acanthaceae)
Wo teige Distr. , Kyadondo Co.,. Kaziy (100° my1Gij.0 059) 7°22
Eee rendOeListh, 2 bUuDuld CO.) LS Of .Katugo, p-l150-m,, GIs
(S6O)iye.0 1.
Kenya. Eastern Prov., Nyeri Distr.,. Mt. Kenya, S ‘slope,
ReGacti, Forest, station, 2200-2300, 2.-3.42.1973, Lb. Ryvarden
C9710, OO), Ll +Iil.
On Asystasia schimperi T. Anders.
NemeMengowWistr., Kyadondo.Co.., > «Namulonge,.. +1200. m,;" “Gy.
CI 4 too) Py ekhabanyvolo, icO0 um, Gi. (205), Tle-Toro iDistre,
Busongora Co., sMweya Lodge, 4930 'm,. Gj....¢149.),. LL.
On Dicliptera sp. (Acanthaceae)
Seber Distrenm Reg. Mt. Elgon, near Mutusyet; 2020121969,
Reis PUVec iL SOM) yl.
On Thunbergia cf. alata Boj. ex Sims. (Acanthaceae)
Kigezi Distr., Rubanda Co., Echuya Forest Reserve, Mile 33
TEomekabare;..2 00,0 -m,.G)6 (237) , 22.
Aeciospores angular, 12-24 x 9-20 pm, wall 0.5-1 pm
thick, irregularly but delicately verrucose, with a few
refracting granules, hyaline. Telia hypophyllous, crust-
like, dark brown. Teliospores varying in shape, cylindric,
clavate or fusiform, truncate or acute at apex, 35-47 x
IZ ioeeAm, ) wall 1-2 pm, vatvapex ‘thickened up to 75.5.) opm,
yellowish, smooth. Pedicel hyaline, persistent.
The specimens reported by Wakefield & Hansford (1949)
as Aecidium acanthacearum Cooke on Justicia spp. were all
placed-with Pp... .thunbergiae by Laundon (1963)... The hosts
mentioned above seem to be new to Uganda, but A. gangetica
and A. schimperi have been reported from Tanzania (Ebbels
& Allen 1979). They also recorded Dicliptera sp., a genus
Seuving. as/a host tor this austin Indiawand Celebes.: i.
alata is type host for P. thunbergiae-alatae P. Henn.,
reported from Namanyonyi in Kyagwe, but the same specimen
was listed from Malawi by Laundon (1963). P. thunbergiae-
alatae has aeciospores with 2-3 pm thick Wa Wl) ueetret tie lie
cf. alata specimen studied by me the wall is hardly above
1 pm thick, thus corresponding well to P. thunbergiae.
Other African Thunbergia species serving as hosts for this
rust are T. atriplicifolia E. Mey., T. natalensis Hook and
536
T. usambarica Lindau.
Puccinia vernoniicola P. Henn. in Engl. Ostafrikan. Pflan-—
zenwelt ip. 150) 111895).
On Vernonia sp. (Asteraceae)
W. Mengo Distr., Mawokota Co., Mpanga Forest, 1150 m, Gj.
9 Boye asl.
Telia hypophyllous, irregular, confluent, compact,
yellow-brown, white when spores germinate. Teliospores
ellipsoid or clavate, lower cell longer than the upper
one, 50-67 x 17-25 pm, wall 1 pm thick, at apex thickened
to 4 pm, hyaline to very pale yellowish, smooth, upper
pore apical, lower near septum. Pedicel hyaline, 40 yum
long, persistent.
This rust, described from Kilimanjaro in Tanzania,
was reported by Wakefield & Hansford (1949) on V. sp. from
Rifle Range in Kiagwe and Kiterera in Busoga. It is also
known from Kenya and S. Africa.
Pucciniosira 'Gissotidis \Wakef.,’ Kew Bull., pi (313, 197.
On Dissotis irvingiana Hook.f. (Melastomataceae)
Mubende Distr., Singo Co., Lwangeta Swamp, W of Kiboga,
hy PARES OG HE ID Lt ak Ts Fis
Telia hypophyllous, cylindrical, in dense groups,
with a peridium. Teliospores in chains, 2-celled, falling
easily apart, ‘ellipsoid, 22-29(-35) (x 12-17 pm, wall 1 pm
thick, hyaline, smooth.
This rust was described by Cooke (1882) as_ both
Aecidium dissotides and Uredo dissotidis. However, the
Sydows (1924) found that it was all the same with the
species described by Wakefield (loc.cit.) on D. incana and
D. sp. from Kyagwe. Later, Wakefield & Hansford (1949),
reported it on D. sp. from Kabaroni in Elgon and from
Kisoro in Kigezi. It is widespread in Africa on Dissotis
spp. and it is also reported on Imbemma sp. from Cameroon
(Heim 1951).
Uredo clerodendrina Vien.-Bourg., Uredineana 5: 231, 1958.
Syn. U. clerodendricola Vien.-Bourg. non P. Henn., Bull.
Soc. ‘Mycol. f2 au sano, 11954.
On Clerodendron myricoides (Hochst.) Vathe (Verbenaceae)
Bugisnu (Distr Won. Bugis 'Co., . Bumesifwa;,.” "S00." my rage
(406) ,..14T1.
Aecia hypophyllous, in small groups, peridium white,
revolute, lacerate. Aeciospores subgloboid, ellipsoid or
ovoid, 22-28)'x% 5-21) ym,) wall 1(-1.5)) pm thick, hyaline,
densely verrucose. Uredinia amphigenous, round, scattered
or in small groups, cinnamon-brown. Urediniospores subglo-
boid, ellipsoid ‘or ‘ovoid, 119-31 x'16=19(-23)' nm, “(wal Deum
thick, sparcely echinulate, and with two equatorial pores
covered with low hyaline papillae, often obscure.
In the original diagnosis only uredinia are de-
scribed. The present material has aecia in close connec-
SOF
tion with the uredinia, and they no doubt belong to the
same species.
U. clerodendrina was described from the Ivory Coast.
Later it is yreportedsaiso,from; Ghana. /°C...myricoides is a
NewWwaPnOSt Lor this wruse which aisenew) to. the. ‘flora: of
Uganda.
Uredovcr wnumbert iia re,’ Budd.) Soc. Bot. Fri: 813.646;
1934,
On Helichrysum cf. gerberifolium Sch. Bip. (Asteraceae)
RaramojgameoLser.., nMatheniko)).Co.,. Mt... sMorimoto, (SE, of
SOGCLONON, 2500KM Ids Oe 19/0, K,A. Lye (M18 By Ole
Uredinia hypophyllous, round, yellow, surrounded by
the brown, cracked epidermis. Urediniospores ellipsoid to
DEGaGgd Obovoid) or? subglobeid;,), 27-32: x 23-27) pm;,)wald,2-2.5
jam thick, very pale yellow, delicately and densely echinu-
late, pores obscure, probably 3-5, scattered.
Bouriquet & Bassino (1965) studying the type speci-
men, found larger urediniospores, 40-42 x 34-36 mpm, with
slightly thicker wall, 3.jm.
This rust is previously known only from Madagascar on
H. vaginatum Humbert.
Uredo hypo€stis-verticillaris n.sp. (Fig. 8)
Uredinia hypophylla, irregularia, dispersa vel in
gregibus parvis, pallide cinnamomea, epidermide longe
obtecta. Urediniosporae subgloboideae, obovoideae, 20-27 x
2 On parece)! 1.5 Mm crasso, hyalino, echinulato,
poris obscuris (duobus equatorialibus instructis?).
Holotype: E. Lang Braun 60a (MHU), Uganda, Karamoja
Distr., Mt. Moroto, 2700-2800 m, 26.02.1960 on Hypoéstes
verticillaris (ise) Soland. ex Roem. & Schult.
(Acanthaceae). Isotype in NPPI.
Laundon (1963) when monographing the rusts on
Acanthaceae, listed only one species with uredinia on
Hypo€stes, viz. Uromyces hypolstis Tarr & Laundon. This
species has larger urediniospores (28-26 x 22-30 wm) with
an amber-coloured, thicker wall 01.5-2.5%5 pm) s,ands 2
approximately equatorial pores. The new species has a
hyaline spore wall which also is more densely echinulate.
Joam
Fig. 8. Uredo hypoestis-verticillaris n.sp.
Urediniospores from type.
538
Uredo isoglossae n. sp. (Fig. 9)
Uredinia hypophylla, irregularia, dispersa, epider-
mide pallide brunnea longe obtecta. Urediniosporae_ sub-
globoideae ad obovoideae vel ellipsoideae, 26-32 x (16-)
21-26 um;, parrete 1-155 nm’ crasso,, hyalino, dense echinge
lato, duobus poris equatorialibus instructis, obscuris.
Holotype:; Li. Ryvarden 19120 (0) 02.-03;°02 519737) $kenyva,
Bastern ‘Prov., Nyeri Distr.,.. Mt. Kenya, Si-slope, vReqacsr
Forest ~“Station;,); 2200-2500° mon’ isoglossa” substrobiriina
C.B. Cl. (Acanthaceae).
Laundon °(1963') mentioned- only the, South) Atrican
species Uredo ecteinanthi Kalchbr. having uredinia on
Isoglossa (Ecteinanthus = Isoglossa), but? “this ruse
species has spores with a yellow-brown and _ sparsely
echinulate wall. Another species, Uredo mitteri (Syd.)
Laundon, described as Coleosporum mitteri Syd. from India
on an Acanthaceae indet., has acc. to the diagnosis
similar but smaller spores than has U. isoglossae which
uredinia are similar to those found in’ Coleosporium.
4 Op
Fig. 9. Uredo isoglossae n-.sp.
Urediniospores from type.
Uredo leucadicola n.sp. (Fig. 10)
Pycnia amphigena, typus 4. Aecia in maculas brunneas,
amphigena, pro maxima parte epiphylla, dispersa in gregi-
bus parvis vel in annulis circa pycnia, peridio albo,
tubiformi. Cellulae peridii plus minusve rectangulares,
pariete externoustriato, 5 pm crasso, “pariete interno,
verrucoso, 3 m crasso. Aeciosporae subgloboideae ad
ellipsoideae vel irregulariae, 22-27 x 16-23 pm, pariete
1.5 pm crasso, hyalino, dense verrucoso. Uredinia amphi-
gena, irregularia, pallide cinnamomea. Urediniosporae
ellipsoideae ad subgloboideae, 22-25 x 18-23 pm, pariete 1
jam crasso, pallide cinnamomeo, echinulato, poris duobus
vel tribus dispersis instructis, obscuris.
Holotype:” HB. .Gjerum)5252° (MHU), §02710.179707 Uganda,
Kagezi, Distr.» Buftunbira. Co.,' iat “the saddles) bemwecn
Muhavusa and Gahinga, 3100 m, on Leucas martinicensis
(Jacgq.) R. Br. (Lamiaceae).
Other specimen seen:
Kenya. Central Prov., Mt. Elgon, Suam Forest Station, 2100
a59
Meee Udo ole, RY Vacdeh "COZ6/7. Oye jail.
Urédinia “and its spores are “much brighter and the
spore wallethicker than “are those of Pi) = leucadis Ps” -& H:
Syd., which also is lacking aecia.
Fig. 10. Uredo leucadicola n.sp.
a. Urediniospores. hb. Aecidiospores.
From type.
redo, paccaeriae PP. & Hs Syd., Ost. boti.°25 (52: 1845°1902.
On Paederia foetens (Hiern) Schum. (Rubiaceae)
Tanzania. Iringa Distr., near Great Ruaha River, 9 km W of
KIGater bridge, 135740970, MeeThulin é- Bo Mhoro. (399)‘ex
NEED. (Ov te
Uredinia hypophyllous, scattered or in groups’ on
reddish to brown leaf spots, sori cinnamon-brown. Ured-
iniospores subgloboid to angular, often triangular or
quadratac in side view, 22-28 x 18-25; pm, wall 1 jm thick,
cinnamon-brown, densely echinulate and with 2-4 scattered
pores.
In the diagnosis the Sydows did not mention pores
while Yen (1970) said they were obscure. In the present
specimen indistinct pores covered by low, hyaline papillae
are recognized in most spores.
Dee peaerrae, described onl. “Mloetidal Lb. Strom, the
Comores, has later been reported scattered on several P.
spp. from India to Ryukyu Islands. It is new to the rust
Plorasot Africa, and P. foetens is a news/hostr.
Ucecompal prgerascumm., Bull “Torrey Bot.. +Club 722 y221;,
1945.
On Inula decipiens E.A. Bruce (Asteraceae)
Mubende Distr., Singo Co., Mile 112 on Kampala-Hoima Road,
near the border of Bunyoro Distr., 1000 m, Gj. (320) II.
Uredinia amphigenous, irregular, long covered by
540
epidermis, with hyaline, thin walled paraphyses, thickened
at apex. Urediniospores obovoid or ellipsoid, 21-27 x 18-
23 pm, wall 1-1.5 pm thick, hyaline or very pale cinnamon-
brown, echinulate, pores obscure.
This species is previously known only from the type
locality,') Viz.) im Kigezi,’.type host isl. | acervatato,
Moore. Cummins (1945) indicated that U. palpigera and some
other similar Uredo spp. probably would be placed in the
genus Phakopsora when telia were found.
Uromyces, jaspilbiicola,Cumm.)), Bull... ‘Torrey Bot.) Clubiea2.
206, 7094s),
On Aspilia africana (Pers.) C.D. Adams. (Asteraceae)
W.). Mengo’ Distr. Busiro Co.,. .Zika: Forest, 160m) wow.
(69),:) ID: Toro Distr," Kyaka) Co. ,) Migenl2 ton ther rodad
Kampala-Fort Portal, W of Nabingora, 1500 m, Gj. (162),
Il; Bunyoro, distr., \Bugahya: Co. , |Hoima,)1500-m, WG. 1o2ceu,
yO ie
Uredinia hypophyllous, cinnamon-brown. Urediniospores
obovoid, sometimes wider than long, 23-30 x 22-30 pm, wall
1.542. (pm thick, in some, spores (slightly thicker sataurne
base, cCcinnamon-brown, delicately echinulate and with 2
equatorial pores.
The rust was described from Kabarmi in the Elgon area
and also mentioned by Wakefield & Hansford (1949). Outside
Uganda | it))is, reported on. A., ‘hotschyi’ from Tanzania ang
Zambia, and from Sudan on Guizotia spp. More doubtful
seems a report on Wedelia sp. from Malawi (cf. e.g. Bisby
& Wiehe 1953). A. africana is a new host for the rust.
Uromyces bidenticolaTArth., Mycologial9? 71, 01 9%s.
Syn. Uredo bidenticola P. Henn., Hedwigia 37: 279, 1898.
On Bidens hirsuta L. (Asteraceae)
W. Mengo Distr., Kyadondo Co., Makerere, 1200 m, Gj. (10),
Me
Uredinia hypophyllous, round, or oblong; brown.
Urediniospores obovoid to ellipsoid, (22-)25-32 x 17-24
pm, wall 1.5 pm thick, cinnamon-brown, echinulate, 2 pores
with low papillae, equatorial or slightly superequatorial.
This rust, widespread in tropical and subtropical
areas, was reported by Wakefield & Hansford (1949) from
Kipayo in Kyagwe, Nkokonjeru in Bugishu and Kabale in
Kigezi. Telia are rarely found in the Eastern Hemisphere
(cf. -Gjerum -1986).
The urediniospore wall thickness is often given as 2-
2.2 pm.
Uromyces blainvilleae Berk., J. Linn. Soc. Bot. Lond. 15:
92 lO hoe
On Blainvillea gayana Cass. (Asteraceae)
Tanzania. ~Ilvinga Distr, ) Ruaha National Park?) 3 (km Neon
Msenbe,.427 «04.1971, Leg. A. Bjornstad:..(.956,.0), .U1 spate.
Uredinia hypophylla, brown. Urediniospores subglo-
541
boid, ellipsoid or somewhat angular, 19-26 x 19-23 pm,
Wall 1 .5-2 pm, , echinulate, cannamon—-brown, with (2-)3(-4)
scattered pores. Telia amphigenous, irregular, often con-
fluent, dark brown. Teliospores subgloboid, 35-38 x 31-37
pm, wall chestnut brown, paler at apex, verrucose, 4.5-5.5
pm thick, at apex thickened to 9 pm, pore apical. Pedicel
hyaline, long, persistent.
This rust has been reported several times from India
and Brazilimainiy ‘on)B. \chomboideae: Cass.) (=B... latifolia
DOw, “but also- on other’ Blainvillea: spp.,') and it Ws also
reported from Galapagos. In Africa it is reported from
Ethiopia, Sudan, Nigeria, and Senegal, form the last
mentioned country on B. gayana (Mayor 1964).
Uromyces hobsonii Vize, Grevillea 4: 115, 1876.
On Jasminum dichotomum Vahl (Oleaceae)
TOLOumuaecr.,) KyakaCo., Mile 127 on the’ Kampala-Fort
Porvalmuoad,, 1500. mm, sG7 010101) LD.
On Jasminum eminii Gilg.
Mubende Distr., Singo Co., Mile 101 on Kampala-Hoima road,
T4000 7 GI. 321) ) yy
On Jasminum meyeri-johannis Engl.
Tanzania. Kilimanjaro, Mwika, 02.01.1960, Bakari B.M. (46)
O+I+III.
Aecia hypophyllous, often making small tumours. Peri-
dium white, revolute. Aeciospores ovoid or angular, 14-22
ed o-toe pity wad Lot Seam chicks! \iverruculose,’') hyaline.
Telia in aecia, dark brown. Teliospores ellipsoid, rarely
Seg ODOs yt \(=55)) 16283 34-37) am, wally 66.5: , jam
chick, at “apex thickened to 9 pm, ‘pale: brown. Pedicel
hyaline, later pale brown, thick-walled.
This rust, described from and widespread in India,
was reported by Wakefield & Hansford (1949) on J.
dichotomum and J. sp. from several localities in Uganda.
Loneasva WLdeldistri bution inwvwAtrica,”) andvin) Asia! it) ).is
also reported from Afghanistan, Nepal and the Philippines.
Js. €Minil is;-a new host for this’ rust.
Uromyces ipomoeae Berk. in Kalchbrenner, Grevillea 11: 19,
1882.
On Ipomoea tenuirostris Choisy (Convolvulaceae)
BugteusdLsccer i... Nie buGiSsu Co... iBumasobo,, 15:00) m, (G7... G40N);,
i fe GP
Uredinia amphigenous, round to elliptical, scattered
or in groups, cinnamon-brown. Urediniospores globoid to
obovoid, 28-33 x 18-27 pm, wall pale cinnamon-brown to
cinnamon-brown, 3-3.5 pm _ thick, echinulate, with 2-4
pores, when 2 equatorial, when 3-4 scattered.
U. ipomoeae was reported by Wakefield (1920) on a
Convolvulaceae, and later by Wakefield & Hansford from
several localities in Uganda, including I. tenuirostris
from Kabale in Kigezi. In Africa it is known from Sudan,
Kenya, Malawi, andS. Africa, and it is also known from
Indonesia.
542
Uromyces melantherae Cooke, Grevillea 10 (1881/82): 127,
1882.
On Melanthera scandens (Schum. & Thonn.) Roberty
(Asteraceae)
Uredinia amphigenous, scattered, cinnamon-brown.
Urediniospores subgloboid, 23-26% VXe O22 3 pm, wall
yellowish brown, 1.5 pm thick, sparcely echinulate with 3
scattered, often obscure pores. Teliospores in the
uredinia, lanceolate oblong to oval, rarely obovoid, 31-45
x 20-24 pm, wall less than 1 fam thick, at apex thickened
to 11 pm, yellowish brown, smooth, pore apical. Pedicel
hyaline, persistent, often longer than the spore.
Doidge (1926) described the urediniospore wall as
about 3 ~m thick, and the teliospores somewhat narrower
(14-18 pm) than seen in the specimen studied by me.
Wakefield): (.1920))° neported’: this )rust \) frome one.
brovnei Sch. Bip.’ “from; Kipayor sand) ont M “ansp ie com
Namanyonyi, both localities in Kyagwe Co. Later Wakefield
& Hansford (1949) added M. pungens Oliv. & Hiern from
Katakwi in Teso to the list. M. scandens is a new host ain
Uganda, but. Mogh~& Hindorf (1971) reported: it. on the ssp.
madagascariensis (Bak.) Wild from Kenya. On M. brownei it
is also reported from Ghana and S. Africa.
Uromyces sasalnsis n.sp. (Fig. 11)
Pycniay samphigena, inygmaculis)foliorum flavispa wd weom
lata, , typus: 4. Aecia amphigena, usque: 250° nm lata dis.
persa vel in annulis circa pycnia, peridio albo, revoluto,
aliquot lacerato. Cellulae peridii rectangulares vel rhom-
biformes:,..30-40 x) 14-16 pm, pariete anterno), 3-4 jJAm crasso,
verruculoso, pariéte jexterno; 3-4) hms) crasso,) “Steiailoe
Aeciosporae subgloboideae vel ellipsoideae, aliquot angu-
lariee, © 14-27 xe 2-207 m,,parietemiet. shim terasso, hyali-
no, dense verruculoso, poris obscuris. Uredinia amphigena,
rotundata vel ovalia, cinnamomea. Urediniosporae ellipsoi-
deae vel globoideae 28-34 x 26-29 pm, pariete. (ty Sa4 pm
crasso, Cinnamomeo dense echinato, spinis fragilibus usque
Spats m longis, 2-4 poris subequatorialibus instructis,
aliquot’ obscuris. ‘Telia hypophylla, retundata vel eltvap—
soidea, dispersa, cinnamomea. Teliosporae obovoideae vel
ellipsoideae, 25-38 x 22-29 pm, pariete 1-1.5 pm crasso,
ad apicem usque 2.5 mm incrassato,, cinnamomeo, laevi, poro
apicali, pedicello hyalino, pariete tenui, collabente,
usque 65 um“longo, sacpe brevi fracto.
Holotype: H.B. Gjerum 376 (MHU), Uganda, Bugisu Distr., N.
Bugisuls Mt se lgon, casa “shut, 32501 m,) -2350n0s to 70 eee
Valeriana volkensii Engl. (Valerianaceae). Isotype in
NEP
Other material:
On Valeriana kilimandscharica Engl.
BugisusDistns, N. Bugis Co.,,: Mt.» Elgon. Sasa Hutu 20mg
GTei(Siip yeas
Two Uromyces spp. are previously described on
Valeriana, VaR Zire Ie valertanae’r (DE. )\) Buck.) vandiaus
valerianae-wallichii Arth. & Cumm. Their urediniospores
543
have thicker walls which are finely echinulate. The telio-
Ssperess of@sthe former “are “smaller than those of -U.
sasalnsis with short pedicel while those in the latter
have scattered, obtuse warts (S. Ahmad 1956).
Fig. 11. Uromyces sasalnsis n.sp.
a. Teliospores. b. Urediniospores.
c. Aeciospores. From type.
544
Host-rust index
In this index are listed the host-rust combinations
published in the papers dealing with the East African rust
fungi (Gjerum 1983, 1984, 1984a, 1985). The page numbers
refer to the fungi.
Abutilon PUuCcCINiawaDwo . dL 4: 245
Acacia Ravenelia hansfordii 4: 263
Acalypha Physopella hansfordii 4: 244
Achyranthes Uredo achyranthicola 4: 264
Aframomum Puccinia aframomi 4: 245
Albizia Ravenelia albiziae-zygiae 4: 263
Ammocharis Aecidium hartwegiae SE ioe)
Andropogon Puccinia eritraeensis Bes 2,
Aneilema Uromyces commelinae 3 fies
Anthephora Puccinia chaseana Wesk2a'6
Anthriscus Puccinia) anthrisecicela 4: 246
Puccinia chaerophylli 4: 247
Artemisia Puccinia artemisiae-afrae SS Sulla
Arthraxon Puccinia arthraxonis-ciliaris 1: 214
Asparagus Puccinia phyllocladiae Scanner
Uredo kabanyoloensis 32 6Syn1S soe
Aspilia Aecidium aspiliae 5 ey, Oo
Uromyces aspiliicola 5: 540
Asystasia Puccinia thunbergiae Sosa
Barleria Puccinia lantanae 5 sa 26
Becium Puccinia becii 53. Zo
Berkheya Puccinia stobaeae var. woodii 5: 533
Bidens Uromyces bidenticola 5s. Ome
Blainvillea Uromyces blainvilleae 531540
Bothriochloa Puccinia duthiae heeuedS
Uromyces clignyi Voapzeo
Brachiaria Physopella africana A Regen eat
Puccinia levis var.
panici-sanguinalis pte ee |
Uromyces setariae-italicae ls 7232
Bromus Puccinia recondita large
Canthium Hemileia canthii 53 514
Carex Puccinia petitiana 22 eo
Uredo caricis-confertae 22 og
Uredo montis-elgonensis 23 pou
Cassia Aecidium torae 4: 240
Ravenelia cassiaecola
var. berkeleyi 43.263
Celosia Aecidium celosiae 4: 238
Cenchrus PucGinia’'cenchri var. africana’ tiiuzls
Cerastium Puccinia arenariae 4:, 247
Chloris Puccinia dietelii eee he:
Uromyces keniensis Ts 5230
Chrysopogon Puccinia duthiae her ig
Clematis Aecidium englerianum 43,238
Coleosporium clematidis 4: 240
Clerodendron Endophyllum superficiale 5s 514
Hemileia scholzii Sean RS
Uredo clerodendrina 5.25536
Clutya
Coffea
Commelina
Conyza
Crassocephalum
Crotalaria
Cyanotis
Cyathula
Cymbopogon
Cynanchum
Cynodon
Cyperus
Deschampsia
Desmodium
Dicliptera
Dicrocephala
Digitaria
Dipcadi
Dissotis
Droguetia
Drymaria
Echinops
Eragrostis
Erlangea
Eriochloa
Eulophia
Euphorbia
Evolvulus
Fimbristylis
Fuirena
Galium
Geranium
Gladiolus
Gossypium
Guizotia
Haplocarpha
Helichrysum
Helictotrichon
Uromyces
Hemileia
cluytiae
vastatrix
Physopella tecta
Uromyces
Aecidium
Aecidium
Puccinia
Uromyces
Uromyces
Puccinia
Puccinia
Puccinia
Aecidium
Puccinia
Puccinia
Puccinia
Puccinia
Puccinia
Uromyces
Puccinia
commelinae
macowanianum
gynurae
crassocephali
crotalariae
commelinae
cyathulicola
andropogonis
nakanishikii
cynanchi
cynodontis
conclusa
cyperi-cristati
cyperi-tagetiformis
morotoensis
notabilis
brachypodii
var. arrhenatheri
Uredo desmodii-ramosissimi
Puccinia
Aecidium
Puccinia
Puccinia
Uromyces
Uromyces
thunbergiae
dichrocephalae
digitariae-velutinae
oahuensis
pegleriae
dipcadi
BUCCAMMOS Maan GaUuSSOLLars
Puccinia
Puccinia
Puccinia
fusispora
leptosperma
dummeri
Physopella hiratsukae
Uredo sp.
Physopella wiehei
Puccinia
Uromyces
Uromyces
erlangeae
setariae-italicae
eulophiae
Melampsora euphorbiae
Uromyces
Puccinia
Puccinia
Puccinia
euphorbiae
desertorum
fimbristylidis
cyperi-tagetiformis
Uredo fuirenae-strictae
Puccinia
Puccinia
Uromyces
Uromyces
difformis
leveillei
geranii
transversalis
Phakopsora gossypii
Puccinia
Aecidium
Aecidium
quizotiae
haplocarphae
echuyaénsis
Physopella nyasalandica
Puccinia
Uredo cf.
Puccinia
kalchbrenneri
humbertii
coronata
NMONMONNMN HUH HK SWKEOLOOWW OS
- OU PWE BONN NU SP HKBWH UNH HUE HOW KUunr-
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546
Heteropogon
Hibiscus
Hoslundia
Hyparrhenia
Hypericum
Hyperthelia
Hypo&stes
Impatiens
Imperata
Indigofera
Inula
Ipomoea
Isoglossa
Jasminum
Justicia
Lantana
Leonotis
Leucas
Lippia
Loranthus
Loudetia
Luzula
Medicago
Melanthera
Melinis
Microglossa
Mitracarpus
Monanthotaxis
Murdannia
Ocimum
Oplismenus
Ornithogalum
Oxygonum
Paederia
Panicum
Paspalum
Pavetta
Pelargonium
Pennisetum
Puccinia
Puccinia
Puccinia
Puccinia
versicolor
exilastvar.
garckiana
hoslundiae
Jevablopleyelal
Phakopsora incompleta
Puccinia
Puccinia
Puccinia
Puccinia
Puccinia
Uromyces
Uromyces
Puccinia
Puccinia
PuccinLa
Puccinia
andropogonicola
eritraeensis
eucomi
hyparrheniae
versicolor
clignyi
hyparrheniae
keniensis
andropogonicola
versicolor
isoglossae
Uredo hypodstis-verticillaris
Dietelia impatientis
Puccinia fragosoana
Ravenelia indigoferae
Uromyces indigoferae
Uredo palpigera
Puccinia holosericea
Uromyces ipomoeae
Uredo isoglossae
Uromyces
Puccinia
Puccinia
Puccinia
Puccinia
hobsonii
semiloculata
natalensis
leonotidicola
leucadis
Uredo leucadicola
Puccinia
Aecidium
fpparcola
? cookeanum
Phakopsora loudetiae
Puccinia
Puccinia
Uromyces
Uromyces
Uromyces
Puccinia
Puccinia
Aecidium
Uromyces
Puccinia
Puccinia
Aecidium
Puccinia
loudetiae
obscura
striatus
melantherae
setariae-italicae
aecidiiformis
bakoyana
popowiae
commelinae
ocimi
flaccida
ornithogaleum
polygoni-amphibii
Uredo paederiae
Physopella cameliae
Uromyces
Uromyces
Puccinia
Aecidium
Puccinia
linearis
setariae-italicae
emaculata
flavidum
pelargonii-zonalis
Phakopsora apoda
Puccinia
cenchri var.
africana
[]-- BPO HSH HH Nb WH UWE UBWHHKHKPUHKNHMNHHnne S&H fb SARK KeKKueL-
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226
248
250
525
210
23
240
220
221
220
228
229
201]
213
226
WA)
537
241
229
264
269
Deg
525
541
53.0
541
533
530
Dze
EAS!
536
Sas:
238
23
222
66
20
542
23:2
4d
520
2B9
68
5S
220
66
260
539
212
234
ZeZ
Zo
510
254
210
abs
Pentas
Peucedanum
Phaseolus
Pilea
Plectranthus
Poa
Polygonum
Pseudarthria
Pseudechinolaena
Psychotria
Pycnostachys
Ranunculus
Rhamnus
Rhynchelytrym
Rhynchosia
Rhynchospora
Ricinus
Rubia
Rubus
Ruella
Rumex
Saccharum
Sanicula
Satureja
Schizachyrium
Securinega
Senecio
Setaria
Sida
Smilax
Sorghum
Spermacoce
Spergula
Spilanthes
Sporobolus
Stylosanthes
Tephrosia
Tetradenia
Thalrcetrum
Themeda
Thunbergia
Tragia
Tristemma
Valeriana
Vangueria
Puccinia
Puccinia
Puccinia
Uromyces
Puccinia
Aecidium
Puccinia
Puccinia
Uromyces
Puccinia
Hemileia
Aecidium
Aecidium
pentadis-carneae
pelargonii-elgonensis
peucedani-kerstenii
appendiculatus
fusispora
plectranthi
brachypodii var.
poae-nemoralis
polygonii-amphibii
pseudarthriae
pseudechinolaenae
holstii
pycnostachydis
ranunculacearum
Endophyllum (?) striatosporum
Puccinia
Puccinia
Uromyces
Puccinia
digitariae-velutinae
levis var.
tricholaenae
rhynchosiae
nabugoboensis
Melampsora euphorbiae
Puccinia
rubiicola
Hamaspora engleriana
Hamaspora longissima
Puccinia
Puccinia
Puccinia
Puccinia
Puccinia
Uromyces
Aecidium
Aecidium
Puccinia
Puccinia
ruelliae
obscuripora
melanocephala
saniculae
menthae
clignyi
fluggeae
dubiosum
hansfordiana
hedbergii
Phakopsora setariae
Uromyces
Puccinia
Puccinia
Puccinia
Puccinia
Puccinia
Puccinia
setariae-italicae
heterospora
smilacis-kraussianae
nakanishikii
lateritia
arenariae
africana
Uredo kabaleensis
Uredo sporoboli-pyramidalidis
Uromyces
tenuicutis
Uredo stylosanthis
Ravenelia tephrosiae
Pucci mira
Uromyces
tetradeniae
wiehei
Phakopsora incompleta
Uromyces
Puccinia
Puccinia
Puccinia
Uromyces
Hemileia
Caagniy.s:
thunbergiae
tragiae
necopina
sasa¥nsis
woodii
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254
207
266
249
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260
269
225
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239
514
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222
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243
53
241
242
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2o2
223
261
530
228
239
509
524
524
Zan
252
250
67
PAPAS
528
247
sii is:
Pea |
22a
253
265
264
Spo NS
Pad |
210
228
D393
262
530
542
516
548
Vernonia
Vigna
Viola
Wissadula
Xyris
Zea
Zehnera
Zornia
Aecidium vernoniae-podocomae
Phakopsora vernoniae
Puccinia vernoniicola
Phakopsora pachyrhizi
Uromyces vignae
Puccinia violae
Puccinia abutili
Uredo xyridis
Puccinia polysora
Puccinia sorghi
Puccinia cephalandrae
Puccinia zorniae
Poe Hw eS
5S
516
536
244
270
262
245
68
224
226
520
262
549
References
Ahmad, S. 1956. Uredinales of West Pakistan. Biologia 2:
26-101.
Arthur, jiJ.C.) 1934... Manual jof\ the rusts: in: United, States
and Canada. Purdue Res. Foundation, Lafayette,
Indiana.
Bisby, GB. | & Wiehe). 0.) 1953). The, rusts: of Nyasaland.
Mycol. Pap. No. 54.
Bouriquet, G. & Bassino, J.-P..1965., Les Urédinées de
Madagascar. Prodrome fl. mycol. Madagascar. V.
Cooke, M.C. 1882.:) Exotic fungi., ,Grevillea 10( 1881-82) :
121-130.
Cummins, G.B. 1945. Descriptions of tropical rusts - VII.
Bul Torreyieet. Chub 723° 205-222.
-~ 1952. Uredinales from various regions. Bull. Torrey
Bor. Club /9:. 212-234.
Dale, W.T. 1955. New species of Uredinales from Trinidad.
MYVCOL. Pap. NO. 159.
Doidge, E.M. 1926. A preliminary study of the South Afri-
Canirust tunga. Bothatia 2s (228.
-e94s.)) SOULK Atrican inust fungi.) Part. Vio (Bothalia)) A's
895-918, 1948.
Ebbels)' Dob. . 1972.) Additions: to ithe mycofilora \of;south-
westernJUcangacs, 3.) JE. Aibrical nat. /, Hist. Soc. Nat,
BANS oy) Ow) SS
- & Allen, D.J. 1979. A supplementary and annotated
list of plant diseases, pathogens and associated
fungi in Tanzania. Phytopath. Pap. No. 22.
Ebon, DOS. 1977) Aecidium mitracarpi Syd.,. a synonym of
Puccinia bakoyana Pat.) & Har.’ Trans. Br. mycol. Soc.
693. 136-137.
Hise. Motes Ban Kleiner’ Beitrag izur)Pilztlora,) won
Tanganyika. Sydowia 25 (1971): 128-129.
Gj@erum, H.B. 1983. East African rusts (Uredinales), mainly
from Uganda. 1. On Poaceae. Mycotaxon 18: 209-234.
- 1984. 2. On Cyperaceae. Mycotaxon 20: 53-63.
- 1984a. 3. On Amaryllidaceae, Commelinaceae, Irida-
ceae, Juncaceae, Liliaceae, Orchidaceae and Xyrida-
ceae. Mycotaxon 20: 65-72.
- 1985. 4. On families belonging to Apetalae and Poly-
petalae. Mycotaxon 24: 237-273.
- 1986. Rust fungi (Uredinales) from Cape Verde
Islands. Bot. Macar. 12-13(1984): 123-138.
Henderson, D.M. 1970... Rust from East Africa. Notes R. Bot.
Gdn Edinb. 30: 395-407.
= ieee RUS Ung i prom mest WALricacs TT. Noes ReuBoOL.
Gdn Edinb. 31: 441-446.
Hughes,» S.J. 1952. Fungi from the’ Gold Coast - 1.) Mycol
Pap. No. 48.
HyJander)) N., )dgerstad, (1. & Nannfelat), J:aA.\1953..) faume-
ratio uredinearum scandinavicarum. Opera Botanica,
ae eee
Jerstad, I. 1956. Reliquiae Lagerheimianae. African Uredi-
Males. Ark, Bet. Ser. 2))( Bay Oo nn pl tty 5OS—B98.
550
Laundon, G.F\:. 1963. Rust fungi 1: On Acanthaceaes Mycol:
Pap stiNOi noo.
Majewski, T. & Nowak, K.A. 1982. Les champignons parasites
recoltés au Rwanda. Bull. "Soc. Myc.) Fr. 9827 369-27.
Mayor, E. 1964. A propos de trois Urédinales du Sénégal.
Build. Soc. Mycol. Fry S03). 828-359.
Mogh,' iM. & Hindorf, H. 1971. Parasitic fungiycollecteasa
Kenya. Nova Hedwigia 21: 479-504.
Nattrass, RM. 1961.‘ Host: list of Kenya) fungi and bacee-
ria. Mycol. Paps (No. 73d".
Nevodovsky, G.S. 1956. Flora sporovykh rastenii Kazakh-
stana. Rzhavtchinnye griby. Alma-Ata.
Petrak, F. 1959. Ein kleiner Beitrag zur, Pilztloramvon
Tanganyika.| Sydowia 13: 223-234.
Stewart, R.B. & Yirgou, D. 1867. indexvotvplant, diseases
in’ Ethiopia." Exp.) "Sta. “Bull Now130,7 Cole. AGrtca,
Haille Selassie I Univ.
Sydow, P. & H. 1924. Monographia Uredinearum IV. Uredineae
imperfectae. Lipsiae.
Viennot-Bourgin, G. 1953. Urédinales d'Afrique (3e note).
Urédinales de Céte d'Ivoire (2e note). Uredineana 4:
VW25=228).
- 1958. Urédinales d'Afrique (5e note). Urédinales de
CSte d'Ivoire (4e note). Uredineana 5: 137-248.
Wakefield, E.M. 1920. Fungi exotici: XXV. Notes on Uganda
fungi. Bull. Misc. Int. R. Bot. Gdns Kew No. 9: 289-
3800.
—) H&o Hansford ieCle.? 1 949 SContribueion towards the
fungus flora of Uganda. IX. The Uredinales of Uganda.
Proc. Linn. Soc bond, Sess. | 164), 1948- 497 pu.
162-198.
Yen, J.-M. 1970. Etude sur les Champignons parasites du
Sud-Est asiatique. XIII: Quelques especes d'Urédinées
de Malaisie. Revue Mycol. 34: 299-339.
Erratum
In No. 3 of this series of papers on East “African
rusts (Mycotaxon 20: 65-72, 1984) on p. 68 Asparagus
officinalis L. is given as a host for Uredo kabanyoloensis
Gj. The host should be A. plumosus Baker.
Page 558 line 2
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MYCOTAXON
VO Pex AV IT ppa Sol-554 October-December 1986
CONFIRMATION OF ASCOSPHAERA APIS IN GEORGIA
R. T. Hanlin and R. A. Saunders
Department of Plant Pathology
Universr.ty of Georgia
Athens, Georgia 30602
ABSTRACT
Ascosphaera apis, cause of chalk brood disease
of bees, is described from honey bee larvae in
Georgia for the first time. Although this is the
first published report of the disease in the state,
it apparently has been known to bee keepers since
the 1920's.
Keywords: Apis mellifera, honey bee, insect
mycosis.
INTRODUCTION
Chalk brood of bees, caused by AScosphaera apss
(Maassen ex Claussen) Olive & Spiltoir, can be a serious
disease of domesticated honey bees (Bailey 1981). During
the past few years several publications (Hitchcock and
Christensen 1972; Thomas and Luce 1972; Christensen
and Gilliam, 1983; Rose et al. 1984) have described
collections of honey bees (Apis mellifera L.), infected
with chalk brood disease from various areas of the United
States. Georgia is not among the areas listed, nor is
the fungus included in published lists of Georgia
ascomycetes (Miller 1941; Hanlin 1963). A search of the
literature also failed to yield any references to the
fungus in Georgia. A mail survey of bee inspectors in
the United States, however, revealed that the disease is
present in Georgia and 32 other states (Menapace and
Wilson 1976). The purpose of this paper is to confirm
the occurrence and identity of A. apis in Georgia.
3
OBSERVATIONS AND DISCUSSION
In April, 1986, specimens of diseased honey bee
larvae were collected from a hive in Madison County,
Georgia and taken to the laboratory for examination. The
larvae were covered with a dense layer of hyphae and
brown spheres (Fig. 1). Examination of these structures
revealed that they were spore cysts [(54)-75-(110) um
diam](Fig. 2) containing spore balls [(12)-16-(26) um
diam](Fig. 3). The spore balls (Fig. 4) were composed of
large numbers of hyaline, one-celled, ellipsoidal spores
[(3.2)-3.5-(4.0) X (1.6)-1.9-(2.0)um] (Fig. 5) that were
straight to slightly curved, with rounded ends. On the
basis of these characteristics, the fungus was identified
as Ascosphaera apis (Rose et al. 1984).
The first report of chalk brood in the United States
was on wild bees collected in Utah in 1965 (Baker and
Torchio 1968). The disease was first found on honey bees
in Tehama County, California in 1968 (Thomas and Poinar
1973), and it was found in numerous other counties in
subsequent years (Thomas and Luce 1972). By 1975 the
disease had been recorded in 33 of the 48 contiguous
states. Whether the disease spread rapidly or had simply
gone unreported is unknown, but the latter seems likely.
In Georgia, e.g., bee keepers reported seeing the disease
in the 1920's (Menapace and Wilson 1976), forty years
before its 'discovery' in the United States, yet there
appears to be no record of its occurrence in the
literature. This emphasizes the importance of publishing
distribution records of fungi, especially those of
economic importance. As of 1975 the disease was still
unrecorded from Louisiana, Mississippi, South Carolina
and Tennessee in the southeastern United States, but it
is probable that it has either been overlooked or ignored.
ACKNOWLEDGEMENT
We thank Martha Christensen and Rodney Roberts for
review of the manuscript.
555
Figures 1-5. Adscosphaera apis. Fig. 1. Honey bee larva
covered with spore cysts. X12. Fig. 2. Spore cysts from
bee larva. X256. Fig. 3. Broken spore cyst with spore
DALle wins 29s) Page 4. Spore! balls. X1450.) Fig.) 5.
Ascospores. X3500.
554
LITERATURE CITED
Bailey, L. 1981. Honey Bee Pathology. Academic Press,
New York.
Baker, G. M., and P. F. Torchio. 1968. New records of
Ascosphaera apis from North America. Mycologia 60:
189-190.
Christensen, M., and M. Gilliam. 1983. Notes on the
Ascosphaera species inciting chalkbrood in honey
bees. Apidologie 14: 291-297.
Hanlin, R. T. 1963. A revision of the Ascomycetes of
Georgia. Univ. Georgia Coll. Agric. Mimeo. Ser. N.
S.0') 17529168;
Hitchcock, J. D., and M. Christensen. 1972. Chalk brood
disease of honey bees in the United States. Amer.
Bee J. 112: 248-249, 254.
Menapace, D., and W. T. Wilson. 1976. The spread of
chalk brood in the North American honey bee, Apis
mellifera. Amer. Bee J. 116: 570-573.
Miller, J. H. 1941. The Ascomycetes of Georgia. Plt.
Dis. Reptr. Suppk. 131: 31-93.
Rose; J. B. 4. My. Christensen, 'and W.. Ts Wilson.1984.
Ascosphaera species inciting chalkbrood in North
America and a taxonomic key. Mycotaxon 19: 41-55.
Thomas, G. M., and A. Luce. 1972. An epizootic of chalk
brood, Ascosphaera apis (Massen [sic] ex Claussen)
Olive and Spiltoir, in the honey bee, Apis mellifera
L. in California. Amer. Bee J. 112: 88-90.
Thomas, G. Ms, andG...0.) Poinar, dr. /1973.4 Reportaor
diagnosis of diseased insects, 1962-1972. Hilgardia
42's 264=359%
MYCOTAXON
VOI AXVIT pp. 555-562 October-December 1986
RECOLTES DE BISCOGNIAUXIA DENNISII
(PYRENOMYCETES, ASCOMYCETES)
DANS LE SUD-QUEST DE LA FRANCE
FRANGOISE CANDOUSSAU *
22, rue Hod-Paris
64000 Pau, France
Résumé: Biscogniauxia dennisii, conmnue jusqu'a présent de
deux stations en Tchécoslovaquie sur Acer et Ulmus, est une
espéce assez fréquente sur Quercus dans trois départements du
Sud-Ouest de la France. Cette espéce n'ayant pas été illustrée
dans la littérature, il nous a semblé utile de le faire a l'aide des
photos. Une description d'aprés la récolte d'Amou (Landes) et
une discussion sur le genre Biscogniauxia sont faites.
Summary :Biscogniauxia dennisii, which has been reported from
two locations in Czekoslovakia on Acer and Ulmus is frequently
encountered on Quercus in three departments in Southwest
France. This species has not been illustrated in the literature. It
seemed useful to present photographs of material. A description
based on a collection from Amou (Landes) and a discussion of
the genus Biscogniauxia is presented.
* Correspondante scientifique de I'herbier de pathologie végétale de I'Université
de Cornell, Ithaca, N. Y. 14853.
556
Biscogniauxia dennisii (Z. Pouzar) Z. Pouzar, Ceska
Mykologie 33:216. 1979
Basionym: Nummularia dennisii Z. Pouzar, Kew Bull. 31:
653. 1976
Stromas crateriformes, erumpents, enfoncés dans Il'écorce du
bois, circulaires a elliptiques, le plus souvent légérement
concaves, 5 - 18 x 4- 8 mm de large et 3 - 4 mm d’é€paisseur.
Marge lisse, surélevée, noire, plus ou moins réguliére, ostioles
proéminentes papillees (Figs. 1, 3). Les stromas en coupe font
apparaitre les péritheces monostiques noirs et brillants, allongés
cylindriques, remplissant de bas en haut presque toute la hauteur
du stroma, et mesurant jusqu'a 3 mm de haut et 0.5 a 1 mm de
large (Figs. 2, 4). La base plus claire du stroma consiste en
matiéres décomposées du bois du support mélangées aux
hyphes du pyrenomycéte (Fig. 2). Ectostroma délimité par une
couche mince carbonacée. Asques 110 - 150 x 14-16 um
courtement stipités, (Figs. 5, 6), appareil apical distinctement
amyloide (Figs. 7, 8). AScospores huit, unisériées, lisses, brun
foncé, Opaques sous le microscope, largement ellipsoides a
citriformes, 14.5 - 18 x 11 - 15 um, la plupart avec bulles de "De
Bary” et sillon germinal visible droit ou legerement en spirale (Fig.
7). Paraphyses cylindriques simples, septees, plus longues que
les asques, 3.5 - 5 um de large (Figs. 5, 6).
Récoltes examinées: FRANCE: 13.ll. 1971 - sur Quercus
pedunculata, Bois de Saint Maur, 32 Mirande, Gers, F.
Candoussau. XI. 1974 - sur Quercus pedunculata, Pouylebon, 32
Mirande, Gers, F. Candoussau. 20. Il. 1983 - sur Quercus
pedunculata, Bois de Garlin (35 km Nord de Pau), Pyrénées
Atlantiques, F. Candoussau, J. Vivant, G. Gilles. 2.lll. 1986 - sur
Quercus pedunculata, Bois d'Amou, Landes, F. Candoussau & G.
Gilles.
Notre récolte concorde bien avec la description de la récolte type
de Pouzar (1976). Les stromas étant d'une grandeur tres variable
(Pouzar,1986) nous ne nous attarderons pas sur les dimensions
plus petites de celles de nos récoltes.
Jusqu'a présent cette espéce a été signalée sur Acer (Slovaquie;
Pouzar, 1976) et sur Ulmus (Bratislava; Pouzar, 1986). Dans le
Soe
Sud Ouest de la France, depuis 1971 (récoltes mal identifiées)
nous ne l'avons récoltée que sur Quercus et elle ne semble
pas rare comme en Tchécoslovaquie ou deux stations sont
seulement connues. La France et la Tchécoslovaquie sont les
deux seuls pays cités jusqu'a présent dans la littérature pour sa
répartition géographique.
Le genre Biscogniauxia O. Kuntze a été émendé par Pouzar en
1979 pour remplacer le genre Nummularia Tul., invalide, car
preoccupe par un genre d'Angiosperme (Nummularia Hill;
Pouzar, 1979).
D'abord, Miller (1928) a rejeté la séparation du genre
Nummularia d'avec le genre Hypoxylon Bull., mais plus tard il a
émendeé le genre Nummularia et y a versé les espéces cupulées,
choisissant N. discreta (Schw.) Tul. comme lectotype du genre
(Miller, 1932).
D'autres auteurs ont suivi, considérant ces espéces distinctes et
differentes du genre Hypoxylon (Jong & Benjamin, 1971).
Eckblad & Granmo (1978) ont cree Nummulariella Eckblad &
Granmo pour les especes cupulées et Pouzar (1979) en
émendant I'ancien genre Biscogniauxia a réuni sous ce nom les
especes cupulées et applaties.
Les espéces du genre Biscogniauxia sont caractérisées
macroscopiquement par des stromas cupulés et des péritheces
se développant dans la base de I'entostroma et atteignant la
surface du stroma par des ostioles plus ou moins allongées, alors
que dans le genre Hypoxylon les péritheces sont situes dans la
partie supérieure de l'entostroma, de plus la base du stroma se
mélange aux matieres décomposées de la plante.
Microscopiquement l'absence d'exospore hyaline des spores,
l'absence de pigment clair soluble dans le KOH, le foramen plat
des asques et l'appareil apical de l'asque, amyloide d'une
maniére diffuse, caractérisent les especes de ce genre.
Récemment Callan & Rogers (1986), apres avoir cultivé
Biscogniauxia marginata (Fr.) Z. Pouzar, B. dennisii et B. repanda
(Fr.) O. Kuntze ont constaté que l'anamorph etait un
Nodulisporium comme dans beaucoup d'autres especes du
genre Hypoxylon et sont arrivés a la conclusion que les deux
genres doivent étre tres proches.
Toutefois les genres Biscogniauxia et Hypoxylon sont deux
genres bien différents et se distinguent particulierement par
l'ontogénie de leur stroma.
Figs. 1 - 2. Biscogniauxia dennisii. - Fig. 1. Stroma sur bois
décortiqué légérement concave; 11x. - Fig. 2. méme stroma, en
coupe faisant apparaitre les péritheces monostiques; 18x.
Figs. 3 - 4. Biscogniauxia dennisii. - Fig. 3. Stroma étalé sur bois
non €corcé; 11x. - Fig. 4. méme stroma faisant apparaitre les
péritheces monostiques; 11x.
Bet ERRATA
560
FANE,
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Figs. 5 - 6. Biscogniauxia dennisii. - Fig. 5. Préparation dans
l'eau. Asques, ascospores et paraphyses; 300x. - Fig. 6. Détail de
la méme préparation mettant en évidence la structure apicale des
asques; 600x.
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Remerciements
Nous remercions Bruno Erb (Erlinsbach, SUISSE) pour les
photographies, le Dr. L. Petrini (Zurich, SUISSE) pour son aide
bibliographique et ses commentaires. Toute notre
reconnaissance au Professeur J. D. Rogers (Washington State
University, Pullman, Wa., U.S.A.) pour I'identification de notre
recolte (Garlin 1983) et pour la révision de notre manuscrit.
Bibliographie
Callan, B. E. & J. D. Rogers (1986). Cultural characters and
anamorphs of Biscogniauxia (= Nummularia) marginata,
B. dennisii and B. repanda. Can. J. Bot. 64: 842 - 847.
Eckblad, F. E. & A. Granmo (1978). The genus Nummularia
(Ascomycetes) in Norway. Norw. J. Bot. 25: 69 - 75.
Jong, S. C. & C. R. Benjamin (1971). North American species of
Nummularia. Mycologia 63: 862 - 876.
Miller, J. H. (1928). Biologic studies in the Sphaeriales Il.
Mycologia 20: 205 - 339.
Miller, J. H. (1932). British Xylariaceae Il. Trans. Br. mycol. Soc.
17 129\-aoo:
Pouzar, Z. (1976). Nummularia dennisii, a new species of
Xylariaceae from Central Europe. Kew Bull. 31: 653 -
655:
Pouzar, Z. (1979). Notes on taxonomy and nomenclature of
Nummularia (Pyrenomycetes). Ceska Mycologie 33: 207
- 219.
Pouzar, Z. (1986). A key and conspectus of Central European
species of Biscogniauxia and Obolarina
(Pyrenomycetes). Ceska Mycologie 40: 1 - 10.
MYCOTAXON
Voie Oe pp 03-620 October-December 1986
NEW SPECIES OF THE LICHEN GENUS XANTHOPARMELIA FROM
SOUTHERN AFRICA (ASCOMYCOTINA: PARMELIACEAE)*
Mason E. Hale, Jr.
Department of Botany, Smithsonian Institution
Washington, DC 20560, U.S.A.
Abstract. Sixty-two new species of Xanthoparmelia are
described from southern Africa: X. acrita Knox & Hale, X. africana
Hale, X. aliphatica Hale, X. barklyensis Hale, X. calvinia Hale,
X. capensis Hale, X. ceresensis Hale, X. competita Hale, X.
crassilobata Hale, X. denudata Hale, X. diacida Hale, X. duplicata
Hale, X. effigurata Hale, X. endochromatica Hale, X. enteroxantha
Hale, X. epigaea Hale, X. equalis Hale, X. esterhuyseniae Hale,
X. evernica Hale, X. globisidiosa Hale, X. gyrophorica Hale, X.
hybrida Hale, X. indumenica Hale, X. karooensis Hale, X. lagune-
bergensis Hale, X. lesothoensis Hale, X. lividica Hale, X. lobuli-
fera Hale, X. minuta Knox & Hale, X. mollis Hale, X. namakwa Hale,
X. namaquensis Hale, X. neosynestia Hale, X. neotasmanica Hale,
X. nuwarensis Hale, X. ochropulchra Hale, X. olifantensis Hale,
X. olivetorica Hale, X. oribensis Hale, X. probarbellata Hale,
X. pustulifera Hale, Nash & Elix, X. rubromedulla Hale, X. sani-
ensis Hale, X. serusiauxii Hale, X. simulans Hale, X. skyrinifera
Hale, X. stenosporonica Hale, X. subcolorata Hale, X. subcrustosa
Hale, X. subnigra Hale, X. subochracea Hale, X. subruginosa Hale,
X. surrogata Hale, X. tablensis Hale, Nash & Elix, X. terricola
Hale, Nash & Elix, X. thamnolica Hale, X. toninioides Hale,
X. transvaalensis Hale, Nash & Elix, X. treurensis Hale, Nash &
Elix, X. tumidosa Hale, X. viridis Hale, and X. wesselsii Hale.
Two new combinations are also made: X. ianthina (Brusse) Hale and
X. lecanoracea (Mull. Arg.) Hale.
*This work was supported by a grant from the Scholarly Studies
Program, Smithsonian Institution. I would like to thank Dirk Wessels,
University of the North, Pietersburg, for arranging forays in
S.W.A./Namibia, eastern Transvaal and the Cape Town area for the
International Association for Lichenology, making possible collecting in
areas not previously visited by lichenologists. I am especially
indebted to Anita Johnson and Dr. C. F. Culberson for determining the
chemistry of several critical species. Dr. J. A. Elix, Dr. David Knox,
Dr. T. H. Nash and Jen Johnston contributed specimens and/or information
on nine of the species included here and valuable loans were received
from Dr. Ove Almborn (LD), Frank Brusse (PRE) and Dr. Harrie Sipman (B).
564
Xanthoparmelia acrita Knox and Hale, sp. nov. Pig. l
Thallus laxe adnatus, terricola vel saxicola, 5-8 cm latus, lobis
sublinearibus, 1-2 mm latis, superne plus minusve maculatus, isidiis
sorediisque destitutus, subtus subconvolutus, pallide brunneus, dense
rhizinosus.
Thallus loosely adnate on rocks, soil, pebbles or soil over rocks,
5-8 cm broad, somewhat darkish greenish yellow; lobes sublinear, quite
variable, 1-2 mm wide, separate to subimbricate, subdivaricately
branched; upper surface weakly to moderately maculate, shiny; isidia and
soredia absent; lower surface weakly to distinctly convolute, without a
distinct paler rim below along the margins, pale brown, moderately to
densely rhizinate, the rhizines fine, brown or blackening, especially at
lobe tips, 0.5-1 mm long, simple to furcate.
Pycnidia rare; conidia bifusiform, 0.6 X 5-6 ym. Apothecia rare,
adnate to substipitate, 2-5 mm in diameter; hymenium 45-48 mm; spores
poorly developed, 6 X 9 pm.
Chemistry: Fumarprotocetraric and usnic acids, with or without
succinprotocetraric acid (both present in holotype).
Type collection: On soil among low sandstone ledges in Fynbos, Ceres
Nature Reserve at Michells Pass, about 1.5 km SW of Ceres on Hwy R46,
elev. ca 600 m, Cape Province, S. Africa, Grid 3319 AD, M. E. Hale 72081,
24 Jan. 1986 (US, holotype; PRE, isotype).
This species is distinguished by the weakly convoluted lobes and
chemistry. It is rather common on open sandstone outcrops and nearby
sandy soil in the karoo and Fynbos of southwestern Cape Province. Some
specimens contain only fumarprotocetraric acid.
Additional specimens examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 72014, 72022, 72024 (US); Karoo Botanical
Garden, Worcester, Grid 3319 CC, Hale 74021, 74022, 74024, 74049, 74051,
74052, 74108 (US), Nash 23494 (ASU, US); near Joubertina, Grid 3323 DD,
Esterhuysen 16596, 16624 (BOL); Tafelberg, Cedarberg, Grid 3219 AC,
Schelpe 1933, 1940 (BOL, US); 32.7 km ESE of Citrusdal, Grid 3219 CA,
Hale 72018, 72015, 72021, 72023, 72115 (US); Witteberg, Grid 3318 DB,
Leighton 263 (BOL, US); Klein Roggeveldberge, 47 km SW of Sutherland,
Grid 3220 DC, Leighton 3150 (BOL, US); Sir Lowry Pass, Grid 3418 BB,
Guthrie & Page s.n. (BOL, US); Mountains above Tulbagh Waterfall, Grid
3318 BD, Stephens s.n. (BOL, US); 23 km SW Ladismith, Grid 3321 CA, Hale
74020 (US); Uniondalespoort, Grid 3323 CA, Hale 730038, 73008 (US).
Xanthoparmelia africana Hale, sp. nov. Fig. 2
Thallus laxe adnatus, saxicola, 10-20 cm latus, lobis sublinearibus,
1.5-5 mm latis, superne vix maculatus, isidiis sorediisque destitutus,
subtus planus, nigricans, dense rhizinosus.
Thallus loosely adnate on rock, firm, sometimes forming thick
pulvinate colonies 10-20 or more cm broad, bright yellowish green; lobes
sublinear, dichotomously branched, 1.5-5 mm wide, crowded and imbricate;
upper surface weakly to rather distinctly white maculate with age; isidia
and soredia absent; lower surface plane, black at the center with a broad
brown zone at the margins, densely rhizinate, the rhizines initially
brown but soon blackening, 0.5-1.5 mm long, simple to furcate.
Pycnidia well developed; conidia bifusiform, 0.5 X 6-8 mm.
Apothecia well developed, subpedicellate, 5-10 mm in diameter; hymenium
42-45 pm; spores 6-7 X 9-12 mm.
Chemistry: Salazinic and usnic acids.
Type collection: Sirimon Track, Mt. Kenya, elev. 3300-3400 m, Kenya,
505
PEPE
I
Figures 1-4. Holotypes of Xanthoparmelia: 1, A. acritas’2, XxX. africana:
3, X. aliphatica; 4, X. barklyensis. Scale = mm.
566
F. R. Fosberg and O. M. Mwangangi 49919, 18 March 1968 (US, holotype;
PRE, isotype).
This conspicuous lichen is confined to higher elevations in the
major mountains in Kenya and Uganda and occurs at 700-800 melevationin
southern and eastern Cape Province. I had originally identified the
material as X. tasmanica (Hook. & Tayl.) Hale, a widespread lichen in
Australasia, southern Africa and the Americas, which has no maculae, a
more membranous thallus and sparse to moderate rhizines.
Additional specimens examined. Kenya: Mt. Elgon, Hedberg 933c (UPS,
US); Mt. Kenya, Griffin 28976 (TRT, US), Hedberg 1714 (UPS, US), Schelpe
2659 (BM, US). Tanzania: Kilimanajaro, Hedberg 1393 (UPS, US), Pocs 6934
(US; also issued in Vézda, Lich. Sel. Exs. 1441), Turrall s.n. (BM, US).
Uganda. Distr. Bugisu: Mt. Elgon, Manum 12/1967 (BM, US). S. Africa. Cape
Province: Meiringspoort, Grid 3322 BC, Hale 73141 (US); 33.8 km W of
Dordrecht, Grid 3126 BC, Hale 73140 (US).
Xanthoparmelia aliphatica Hale, sp. nov. Fig. 3
Thallus laxe adnatus, saxicola, 10-12 cm latus, lobis
subirregularibus, 3-6 mm latis, superne emaculatus, isidiis sorediisque
destitutus, subtus planus, niger, dense rhizinosus.
Thallus loosely adnate on rock, 10-12 cm broad, yellowish green;
lobes subirregular, apically subrotund, 3-6 mm wide, contiguous to
imbricate; upper surface continuous, emaculate, rugose and cracked with
age; isidia and soredia absent; lower surface plane, black, densely
rhizinate, the rhizines simple, 0.5-1 mm long.
Pycnidia numerous; conidia bifusiform, 0.6 X 5-7 um. Apothecia
numerous, adnate, 5-8 mm in diameter; hymenium 42-45 um; spores 5-7 X
10-12 mum.
Chemistry: Constipatic and associated fatty acids, usnic acid.
Type collection: Exfoliating granitic outcrops in Karoo, 78.6 km S
of Sutherland on Hwy R354, elev. ca 900 m, Cape Province, S. Africa, Grid
8220 DC, M. E. Hale 73107, 29 Jan. 1986 (US, holotype; PRE, isotype).
This large, conspicuous lichen has a black lower surface, a rare
trait in fatty acid-containing species shared only by X. subnigra Hale
and X. transvaalensis Hale, Nash & Elix (below). It is otherwise
similar to X. barklyensis Hale, also described below and occurring with
it at the type locality, which differs in having a pale brown lower
surface.
Additional specimens examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 73125 (US); 40 km E of Sutherland, Grid
3221 AC, Brusse 3336 (PRE).
Xanthoparmelia barklyensis Hale, sp. nov. Fig. 4
Thallus laxe adnatus, saxicola, 8-15 cm latus, lobis subirregular-
ibus, 2.5-5 mm latis, centro laciniatis, superne emaculatus, isidiis
sorediisque destitutus, subtus planus, pallide brunneus, modice
rhizinosus.
Thallus loosely adnate on rock, 8-15 cm broad, bright yellow green;
lobes subirregular to sublinear, 2.56-5 mm wide, contiguous to imbricate,
the center becoming laciniate; upper surface continuous, shiny,
emaculate; isidia and soredia absent; lower surface plane, pale brown,
moderately rhizinate, the rhizines pale brown, simple, 0.5-1 mm long.
Pycnidia numerous; conidia bifusiform, 0.6 X 5-6 ym. Apothecia
numerous, adnate, 3-5 mm in diameter; hymenium 40-45 um; spores 5-7 X
9-11 mm.
aOW
Chemistry: Constipatic and associated fatty acids, usnic acid.
Type collection: Barkly Pass, about 18 km N of Elliott along Hwy
R58, elev. ca 1800 m, Cape Province, S. Africa, Grid 3127 BB, M. E. Hale
73015, 9 Feb. 1986 (US, holotype; PRE, isotype).
This species appears to be a morph of X. aliphatica Hale with a pale
brown lower surface, occurring mostly in the eastern Cape Province region
and adjacent Orange Free State on Cave sandstone. The only other obvious
relative of this large lichen with a pale lower surface is X._
subdecipiens (Vain.) Hale, which is adnate to tightly adnate.
Additional specimens examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 73001, 73009, 73012, 73013 (US); 78.6 km S
of Sutherland, Grid 3220 DC, Hale 73104 (US); Nieuwoudtville, Grid 3119
AC, Hale 72005 (US); NE of Rouxville, Grid 3026 BD, Hale 74001, 74002
(US); 4.1 km NE of Dordrecht, Grid 3127 AC, Hale 73000 (US). Orange Free
State: Brandwagrots, Golden Gate National Park, Grid 2828 BC, Hale, 74003
(US); 4.8 km west of junction Hwy 56 and 26, near Lady-
brand, Grid 2927 AB, Hale 74048 (US).
Xanthoparmelia calvinia Hale, sp. nov. Fig. 5
Thallus arcte adnatus, saxicola, 4-6 cm latus, lobis subirregular-
ibus, 0.5-2 mm latis, contiguis, superne maculatus, centrum versus valde
rugoso-—isidiatus, isidiis globosis, eruptentibus, subtus planus,
brunneus, modice rhizinosus.
Thallus tightly adnate on rock, 4-6 cm broad, light brownish yellow
green; lobes short, subirregular, 0.5-2 mm wide, contiguous; upper
surface faintly maculate, shiny, becoming strongly rugose and finally
coarsely isidiate at the center, the isidia globose, ca 0.2 mm in
diameter and 0.2-0.3 mm high, pustulate and bursting open to reveal a
hollow center; lower surface plane, brown to dark brown, moderately
rhizinate, the rhizines brown, simple, 0.2-0.4 mm long.
Pycnidia and apothecia lacking.
Chemistry: Three undetermined substances forming gray or brownish
spots on the TLC plates after visualization with sulfuric acid (deter-
mined by C. F. Culberson and A. Johnson), usnic acid.
Type collection: Low dolerite ridges, Akkerendam Nature Reserve near
Calvinia, elev. 1000 m, Cape Province, S. Africa, Grid 3119 BD, M. E.
Hale 75103, 29 Jan. 1986 (US, holotype; PRE, isotype).
This unusual lichen has hollow globose isidia not unlike those of X.
evernica Hale (below). Chemically it is related to X. competita Hale
(below), which lacks isidia. It is known only from eastern Namaqualand.
Xanthoparmelia capensis Hale, sp. nov. Fig. 6
Thallus adnatus, saxicola, 4-6 cm latus, lobis subirregularibus,
1.2-2.0 mm latis, superne emaculatus, isidiatus, isidiis subcylindricis
vel globosis, subtus planus, niger, modice rhizinosus.
Thallus adnate on rock, 4-6 cm broad, pale yellowish green; lobes
subirregular, short, 1.2-2.0 mm wide, contiguous; upper surface contin-
uous, shiny, emaculate, isidiate, isidia subcylindrical to globose,
0.15-0.2 mm in diameter, 0.4-0.7 mm high, breaking off apically; soredia
absent; lower surface plane, black, sparsely to moderately rhizinate, the
rhizines black, simple, about 0.5 mm long.
Pycnidia and apothecia lacking.
Chemistry: Salazinic and usnic acids, chalybaeizans unknown.
Type collection: Low sandstone ledges in pasture along road, 27 km
E of Swellendam on N2, elev. ca 200 m, Cape Province, S. Africa, Grid
568
3420 BA, M. E. Hale 74046, 3 Feb. 1986 (US, holotype; PRE, isotype).
This rare species, widely distributed in southern Cape Province, is
similar to stictic acid-containing X. conspersa (Ach.) Hale in adnation
and lobe configuration but has a different chemistry. Xanthoparmelia
australasica Galloway from Australia and New Zealand, another isidiate
species with salazinic acid (but without the chalybaeizans unknown), is
more loosely adnate and has sublinear lobes.
Additional specimens examined. S. Africa. Cape Province: Road to
Bulhoek, Grid 3126 DC, Hale 74100 (US); Paarl Rock, Grid 3318 DB, Hale
73105 (US); 19.9 km NW Kango Caves, Grid 3322 AC, Hale 73123 (US).
Xanthoparmelia ceresensis Hale, sp. nov. Fig. 7
Thallus adnatus, fragilis, saxicola, 5-7 cm latus, lobis sub-
linearibus, congestis, 0.5-0.7 mm latis, superne continuus vel leviter
reticulato—-maculatus, isidiis sorediisque destitutus, subtus planus,
niger, sparse rhizinosus.
Thallus adnate to loosely adnate on rock and soil over rock,
pulvinate, easily breaking apart, 5-7 cm broad, darkish yellow green;
lobes sublinear, 0.5-0.7 mm wide, sparsely dichotomously branched,
crowded and appressed-imbricate; upper surface continuous or faintly
reticulate-maculate and rugose at the tips, shiny; isidia and soredia
absent; lower surface plane, black, very sparsely rhizinate, the
rhizines black, coarse, 0.3-0.5 mm long.
Pycnidia well developed; conidia bifusiform, 0.5 X 5-7 mpm.
Apothecia not fully developed; spores not found.
Chemistry: Microphyllinic and normicrophyllinic acids (in equal
concentration), 4-O-methylolivetoric acid (trace) and usnic acid
(determined by C. F. Culberson and A. Johnson).
Type collection: Low sandstone ledges in Fynbos, Ceres Nature
Reserve at Michells Pass, 1.5 km SW of Ceres on Hwy R46, elev. ca 600 m,
Cape Province, S. Africa, Grid 3319 AD, M. E. Hale 72,008, 24 Jan. 1986
(US, holotype; PRE, isotype).
This curious species, known only from one locality in Fynbos, forms
brittle, somewhat pulvinate colonies. The appressed sublinear lobes and
faint reticulate-maculation, as well as the unusual chemistry (first
reported occurrence of microphyllinic acid in the genus), are unique.
Xanthoparmelia competita Hale, sp. nov. Fig. 8
Thallus arcte adnatus, saxicola, 3-6 cm latus, lobis subirregular—
ibus, 0.7-1.5 mm latis, contiguis, superne continuus vel leviter
maculatus, centrum versus rugoso-—bullatus, subtus planus, pallide
brunneus, modice rhizinosus.
Thallus tightly adnate on rock, 3-6 cm broad, dull yellowish green;
lobes subirregular, 0.7-1.5 mm wide, short and contiguous; surface
continuous to finely reticulately rugose or maculate at the tips,
becoming strongly rugose-bullate toward the center, isidia and soredia
absent; lower surface plane, pale brown, moderately rhizinate, the
rhizines pale brown, simple, 0.2-0.4 mm long.
Pycnidia common; conidia bifusiform, 0.6 X 6-7 pm. Apothecia
numerous, adnate, 2-4 mm in diameter; hymenium 33-38 pm; spores 5-6 X
9-10 mm.
Chemistry: The same three undetermined substances occurring in X.
calvinia (above), usnic acid (determined by C. F. Culberson and A.
Johnson).
Type collection: Large domes of coarse conglomerate in Fynbos, 3.6
569
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Figures 5-8. Holotypes of Xanthoparmelia: 5, X. calvinia; 6, X.
capensis; 7, X. ceresensis; 8, X. competita. Scale = mm. afi
570
km N of Hwy R62 on Kruisrivier Road, E of Calitzdorp, elev. ca 500 m,
Cape Province, S. Africa, Grid 3321 BB, M. E. Hale 78019, 30 Jan. 1986
(US, holotype; PRE, isotype).
This species is closely related to X. calvinia Hale (above) in
chemistry. While lobe configuration is similar, X. competita is a
brighter yellow and lacks isidia. It appears to be restricted to
southern Cape Province.
Additional specimen examined. S. Africa. Cape Province: 24 km N of
Willowmore, Grid 3322 AB, Hale 73503 (US).
Xanthoparmelia crassilobata Hale, sp. nov. Fig. 9
Thallus laxe adnatus, terricola, coriaceus, 4-8 cm latus, lobis
sublinearibus, 3.5-7 mm latis, superne emaculatus, isidiis sorediisque
destitutus, subtus subconvolutus, pallide brunneus, modice rhizinosus.
Thallus loosely adnate to free growing on soil and loose pebbles,
leathery and firm, bright yellowish green, 4-8 cm broad; lobes broadly
sublinear, 3.5-7 mm wide, separate to imbricate; upper surface con-
tinuous, shiny, emaculate, rugose with age; isidia and soredia absent;
lower surface plane to broadly convoluted, pale brown, rugose, moderately
rhizinate, the rhizines pale brown, 1-1.5 mm long.
Pycnidia not found. Apothecia substipitate, 5-14 mm in diam-
eter, the rim rolled inward; hymenium 43-48 wm; spores 6-7 X 9-11 um.
Chemistry: Salazinic and usnic acids.
Type collection: On soil on flat dolerite outcrops in pasture, 21 km
NW of Sutherland on Hwy R354, elev. 1500 m, Cape Province, S. Africa,
Grid 3220 BC, M. E. Hale 740538, 29 Jan. 1986 (US, holotype; PRE,
isotype).
The thallus of this soil-inhabiting lichen is large, leathery, and
bright yellow. At first it is easily mistaken for an aberrant X.
colorata (Gyelnik) Hale, a@ very common saxicolous lichen in the Karoo,
which has migrated to the soil habitat. However, the lower surface is
clearly brown, not at all blackened, and the only medullary substance is
salazinic acid without norstictic acid. It is much larger, broader lobed
and more coriaceous than X. taractica (Kremplh.) Hale, a soil—inhabiting
or saxicolous lichen in Australasia and South and Central America. It
has been found only in karoo in the south-central part of Cape Province.
Additional specimens examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 74077, 74078, 74079, 74080, 74081, 74082,
74083, 74084 (US); 23 km SW of Ladismith, Grid 3321 CA, Hale 74029 (US);
40 km NW of Merweville, Grid 3221 CA, Brusse 3413 (PRE).
Xanthoparmelia denudata Hale, sp. nov. Fig. 10
Thallus laxe adnatus, saxicola, 5-8 cm latus, lobis sublinearibus,
0.6-1.3 mm latis, apice subadscendentibus, superne valde maculatus,
isidiis sorediisque destitutus, medulla pro parte ochracea, subtus
planus, brunneus, centrum versus ochraceus, sparsissime rhizinosus.
Thallus loosely adnate on rocks, forming continuous mats 5-8 cm
broad or discrete, nearly umbilicate colonies 2-4 cm broad, coalescing
into mats up to 15 cm broad, dark yellowish green; lobes sublinear,
0.6-1.3 mm wide, separate, ascending at the tips; upper surface
distinctly maculate, shiny, transversely cracked with age; isidia and
soredia absent; lower surface plane, rugose, brown but eroding toward the
center to expose the ochraceous medulla, black at the lobe tips, rhizines
very sparse, up to 1 mm long, brown or blackening, coarse.
otk
ed
Figures 9-12. Holotypes of Xanthoparmelia: 9, X. crassilobata; 10, X.
denudata; 11, X. diacida; 12, X. duplicata. Scale = mm.
57
Pycnidia lacking. Apothecia rarely developed, substipitate, 3-6 mm
in diameter; hymenium 45-50 um; spores 6-7 X 8-10 um (data from Hale
73004).
Chemistry: Salazinic and usnic acids, skyrin.
Type collection: Large exposed sandstone ledges in pasture, 1 km S
of Nuwerus on Hwy R363, east of N7, elev. ca 500 m, Cape Province, S.
Africa, Grid 3118 AB, M. E. Hale 72041, 26 Jan. 1986 (US, holotype; PRE,
isotype).
The pale ochraceous orange lower surface is easily recognizable in
the field. The lower cortex erodes away with age to reveal the heavily
pigmented lower medulla. The lobe configuration and nearly bare, rugose
lower surface resembles X. subcolorata Hale (see below), which has a
stronger tendency to form umbilicate thalli, broader lobes (1.5-4 mm
wide) and protocetraric acid. Xanthoparmelia hyporhytida (Hale) Hale,
also restricted to Namaqualand, has salazinic acid and a similar rugose
lower surface eroding with age but lacks skyrin and maculae.
Additional specimens examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 72010 (US); 44 km N of Vanrhynsdorp, Grid
3118 BC, Hale 73004, 74114 (US); 1 km E of Springbok, Grid 2917 DB, Hale
72011 (US).
Xanthoparmelia diacida Hale, sp. nov. Fig. 11
Thallus adnatus, saxicola, 5-8 cm latus, lobis subirregularibus, 2-4
mm latis, superne emaculatus, isidiis sorediisque destitutus, medulla pro
parte pigmentosa, subtus planus, niger, modice rhizinosus.
Thallus adnate to somewhat loosely attached on rock, 5-8 cm broad,
bright yellow green; lobes subirregular, apically rotund, 2-4 mm wide,
imbricate and short lobulate with age; upper surface continuous, shiny or
rarely white pruinose at the tips, emaculate; isidia and soredia absent;
lower part of medulla dull reddish; lower surface plane, black,
moderately rhizinate, the rhizines black, simple, 0.5-1 mm long.
Pycnidia not found. Apothecia adnate, 4-5 mm in diameter; hymenium
45 um; spores 6-7 X 12-15 pm.
Chemistry: Psoromic, protucetraric and usnic acids, an unidentified
fatty acid and the "schenckiana" pigment (determined by C. F. Culberson
and A. Johnson).
Type collection: Sandstone ridge in high veld, 33.8 km W of
Dordrecht on Hwy R56, elev. 800 m, Cape Province, S. Africa, Grid 3126
BC, M. E. Hale 73015, 8 Feb. 1986 (US, holotype; PRE, isotype).
Two other non-isidiate species with psoromic acid, X. huachucensis
(Nash) Egan and X. nigropsoromifera (Nash) Egan, also occur in S.
Africa. They are both smaller, more tightly adnate lichens lacking both
protocetraric acid and the "schenckiana" pigments. Xanthoparmelia
schenckiana (Mill. Arg.) Hale, which contains protocetraric acid and the
same unidentified dull rusty colored pigment (forming a pale yellow
streak near norstictic acid on TLC plates and reacting UV+ yellow
fluorescent), is superficially similar but lacks psoromic acid and is
usually heavily pruinose. This species is confined to the southern end
of the Drakensbergs.
Additional specimen examined. S. Africa. Orange Free State: 18 km E
of Marquard, Grid 2827 CB, Venter 4487 (PRE).
Xanthoparmelia duplicata Hale, sp. nov. Fig. 12
Thallus laxe adnatus, terricola, 3-5 cm latus, lobis sublinearibus,
1.5-4 mm latis, superne emaculatus, isidiis sorediisque destitutus,
Be
medulla pro parte ochracea, subtus convolutus, brunneus, modice
rhizinosus.
Thallus loosely attached to nearly free growing on soil and
pebbles, 3-5 cm broad but coalescing into larger colonies, yellowish
green; lobes sublinear, 1.5-4 mm wide, separate to imbricate, more or
less subascending; upper surface weakly to distinctly maculate, shiny;
isidia and soredia absent; medulla ochre in lower parts; lower surface
weakly convoluted, brown with ochre spots, sparsely rhizinate, the
rhizines brown, 1-2 mm long, coarse, simple to furcate with splayed
tips.
Pycnidia not seen. Apothecia well developed, substipitate, 4-8 mm
in diameter; hymenium 42-45 um; spores 6-7 X 9-12 wm.
Chemistry: Protocetraric, 4-O-demethylbarbatic (trace) and usnic
acids, a new depside related to diffractaic acid and skyrin (determined
by C. F. Culberson and A. Johnson).
Type collection: On soil over quartzite pebbles in flat pasture
land, 19.5 km NNW of Vanrhynsdorp on east side of Hwy N7, elev. 200 m,
Cape Province, S. Africa, Grid 3118 BC, M. E. Hale 73102, 26 Jan. 1986
(US, holotype; PRE, isotype).
This unique soil-inhabiting lichen is alone among the convoluted
species in having this series of acids and the pigment skyrin. It was
collected in the quartzite pebble-paved Knersvlakte region of
Namaqualand along with X. epigaea Hale (below).
Additional specimen examined. S. Africa. Cape Province: 44 km N of
Vanrhynsdorp, Grid 3118 BC, Hale 73106 (US).
Xanthoparmelia effigurata Hale, sp. nov. Fig. 13
Thallus adnatus, coriaceus, 6-12 cm latus, lobis sublinearibus,
brevibus, 1.5-2.5 mm latis, superne valde effigurato—maculatus, isidiis
sorediisque destitutus, subtus planus, niger, sparse rhizinosus.
Thallus adnate to loosely adnate on rock, firm and leathery, pale
yellowish green, 6-12 cm broad; lobes sublinear, 1.5-2.5 mm wide,
dichotomously branched; upper surface strongly effigurate maculate, the
maculae irregularly elongate, sometimes coalescing, shiny, becoming
transversely cracked, somewhat convex with age; isidia and soredia
absent; lower surface plane, black with a brown zone at the tips,
sparsely to moderately rhizinate, the rhizines coarse, brown to black,
0.5-1 mm long.
Pycnidia common; conidia bifusiform, 0.5 X 6-8 uum. Apothecia
commonly developed, substipitate, the rim upturned, 3-7 mm in diameter;
hymenium 40-45 pm; spores 6-7 X 9-11 pm.
Chemistry: Salazinic acid, chalybaeizans unknown, and usnic acid,
with or without traces of norstictic acid.
Type collection: Schist ledges on hillside, Meiringspoort, 18 km N
of De Rust on east side of Hwy R29, elev. 690 m, Cape Province, S.
Africa, Grid 3322 BC, M. E. Hale 73016, 1 Feb. 1986 (US, holotype; PRE,
isotype).
This effigurate—maculate lichen, common on exposed sandstone
outcrops in Namaqualand and southwestern Cape Province, is more or less
related to the X. hypoleia group, and is in fact the only one (with X.
namakwa Hale (below)) containing salazinic acid. It is, however, not as
loosely adnate, almost always being collected with the rock substrate
and having more leathery, separate somewhat convex lobes. Xanthopar-
melia namakwa Hale (below), a morph with a pale brown lower surface,
differs chemically in usually producing salazinic and norstictic acids
in equal concentration, only rarely with the chalybaeizans unknown.
574
Additional specimens examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 73100, 73118 (US); 1 km E of Springbok,
Grid 2917 DB, Hale 72101 (US); Kamieskroon, Grid 3017 BB, Hale 73002,
73127 (US); 1 km S of Nuwerus, Grid 3118 AB, Hale 72094, 72100 (US);
Vanrhyns Pass, Grid 3119 AC, Hale 73130 (US); Nieuwoudtville, Grid 3119
AC, Hale 72006, 72102 (US); Anenouspas, Grid 2917 BA, Hale 73126 (US); 8
km S of Lainsburg, Grid 3320 BB, Hale 75113 (US), Brusse 3535 (LD, PRE);
5 km NW of Wupperthal, Grid 3219 AC, Brusse 3211 (PRE).
Xanthoparmelia endochromatica Hale, sp. nov. Fig. 14
Thallus arcte adnatus, saxicola, 1-2 cm latus, lobis sublinearibus,
0.3-0.9 mm latis, centrum versus opuntioidie constrictis, superne
emaculatus, isidiis sorediisque destitutus, medulla aurantio-ochracea,
subtus planus, niger, sparse rhizinosus.
Thallus tightly adnate on rock, 1-2 cm broad, yellowish green but
blackening with age at the center; lobes sublinear at the margins,
strongly black rimmed, 0.3-0.9 mm wide, contiguous, crowded and becoming
opuntioid-constricted at the center; isidia and soredia absent; medulla
dull rusty red; lower surface plane, black, shiny, sparsely rhizinate,
the rhizines coarse, 0.2-0.3 mm long.
Pycnidia and apothecia lacking.
Chemistry: Gyrophoric and usnic acids, "schenckiana" pigment and
faint unidentified spots.
Type collection: On Table Mountain sandstone ledges, along trail
from Platteklip Gorge to Woodhead Reservoir, Table Mountain Nature
Reserve, elev. 950 m, Cape Province, S. Africa, Grid 3318 CD. M. E. Hale
72081, 23 Jan. 1986 (US, holotype; PRE, isotype).
This inconspicuous lichen known from two high elevation localities
in southwestern Cape Province is characterized by both chemistry and
morphology. It is only the third species of Xanthoparmelia with gyro-
phoric acid as the main component (the other two are X. leucostigma
Brusse and X. gyrophorica Hale (below)). The lobes become rounded and
opuntioid-constricted toward the center of the thallus and appear quite
unlike any other species in the genus. It is intermixed with X.
olivetorica Hale (below), which lacks any pigment and has entirely
sublinear lobes.
Additional specimens examined. S. Africa. Cape Province: Summit of
Swartberg Pass on R328, Grid 3322 AC, Hale 72082, 72117 (mixtures with X.
stenosporonica Hale (below)) (US).
Xanthoparmelia enteroxantha Hale, sp. nov. Fig. 15
Thallus laxe adnatus, saxicola, fragilis, 6-10 cm latus, lobis
sublinearibus, elongatis, 1-1.5 mm latis, superne valde maculatus,
isidiis sorediisque destitutus, medulla omnino pallide salmoneo-
ochracea, subtus planus, niger, modice rhizinosus.
Thallus loosely adnate on rock or on soil over rocks, rather stiff
and fragile, 6-10 cm broad, greenish yellow; lobes sublinear, elongate,
1-1.5 mm wide, dichotomously branched, separate to imbricate; upper
surface clearly maculate, shiny; isidia and soredia absent; medulla
uniformly salmon orange; lower surface plane, dark brown or blackening,
sparsely to moderately rhizinate, the rhizines brown to black, rather
coarse, 0.5-1 mm long, simple to furcate.
Pycnidia and apothecia lacking.
Chemistry: Salazinic and usnic acids, skyrinol, oxyskyrin, skyrin
(trace) and a fourth unidentified anthraquinone (determined by C. F.
575
Figures 13-16. Holotypes of Xanthoparmelia: 13, X. effigurata; 14, Xx.
endochromatica (lower insert X10); 15, X. enteroxantha; 16, X. epigaea.
Scale = mm.
576
Culberson and A. Johnson).
Type collection: Rock outcrops along Hwy R328, ca 22 km NW of Kango
Caves, elev. ca 1000 m, Cape Province, S. Africa, Grid 3822 AC, M. E.
Hale 74044, 31 Jan. 1986 (US, holotype; PRE, isotype).
No other species of Xanthoparmelia, excepting X. endomiltoides
(Nyl.) Hale, X. endochromatica (Hale) (above) and X. ianthina (Brusse)
Hale (comb. nov. based on Parmelia ianthina Brusse in Knox and Brusse,
Jour. S. African Bot. 49: 152. 1983), has a completely pigmented
medulla. This species occurs at higher elevations in southwestern Cape
Province.
Additional specimens examined. S. Africa. Cape Province: Bainskloof,
Grid 3319 CA, Hale 72008 (US); Platteklip Gorge to Woodhead Reservoir,
Table Mountain Nature Reserve, Grid 3318 CD, Hale 72007 (US).
Xanthoparmelia epigaea Hale, sp. nov. Fig. 16
Thallus laxe adnatus, terricola, coriaceus, 3-5 cm latus, lobis
sublinearibus, elongatis, 1.5-5 mm latis, superne emaculatus, isidiis
sorediisque destitutus, subtus convolutus, pallide brunneus, dense
rhizinosus.
Thallus free growing on soil and pebbles, leathery, covering
extensive areas but breaking up into individual colonies 3-5 cm broad,
light yellowish green; lobes sublinear, elongate and little branched,
1.5-5 mm wide; upper surface continuous, shiny, emaculate, rugose with
age; isidia and soredia absent; lower surface moderately to strongly
convoluted, pale brown, densely rhizinate, the rhizines pale brown,
0.5-1.5 mm long, simple to furcate.
Pycnidia common; conidia bifusiform, 0.56 X 6-7 pm. Apothecia
sparsely developed, adnate, 2-4 mm in diameter; hymenium 42-45 pm;
spores 5-6 X 9-10 mm.
Chemistry: Constipatic and associated fatty acids and usnic acid.
Type collection: Quartzite pebbles and schist in flat pasture land,
19.5 km NNW of Vanrhynsdorp on east side of Hwy N7, elev. 200 m, Cape
Province, S. Africa, Grid 3118 BC, M. E. Hale 73020, 26 Jan. 1986 (US,
holotype; PRE, isotype).
This soil-inhabiting lichen, probably endemic to the Knersvlakte
region of Namaqualand, is related to X. chlorochroa (Tuck.) Hale but
differs in being less distinctly convoluted and in having a fatty acid.
All other convoluted species contain K+, P+ depsidones or norlobaridone.
Additional specimen examined. S. Africa. Cape Province: Same
locality as holotype, Hale 73014 (US); Bowesdorp, Grid 3017 BB, Stokoe
7719 (BOL, US).
Xanthoparmelia equalis Hale, sp. nov. Fig. 17
Thallus arcte adnatus, saxicola, mollis, 5-8 cm latus, lobis
sublinearibus, 0.5-1.1 mm latis, congestis vel centro areolato-—bullatis,
superne emaculatus, isidiis sorediisque destitutus, subtus planus,
pallide brunneus, modice rhizinosus.
Thallus adnate to tightly adnate on rocks, soft, 5-8 cm broad,
yellowish green; lobes sublinear, short, 0.5-1.1 mm wide, becoming
transversely cracked, rather crowded and imbricate to areolate-bullate at
the center; upper surface continuous, shiny, faintly reticulate-—
maculate at the tips; isidia and soredia absent; lower surface plane,
pale brown, sparsely to moderately rhizinate, the rhizines pale brown,
0.3-0.5 mm long, simple.
Pycnidia and apothecia lacking.
a7 7
Chemistry: Evernic, lecanoric (trace) and usnic acids (determined by
C. F. Culberson and A. Johnson).
Type collection: Laguneberg Mountains, NE of Mile 72, Distr.
Omaruru, S.W.A./Namibia, Grid 2114 CC, M. E. Hale 75101, 8 Jan. 1986 (US,
holotype; PRE, isotype).
This Namibian endemic is externally close to X. lagunebergensis Hale
(protocetraric acid present) and X. serusiauxii Hale (lecanoric acid
present), which occur at the same locality. It differs from the common X.
lecanoracea (Mull. Arg.) Hale (comb. nov. with basionym Parmelia
lecanoracea Mull. Arg., Flora 71: 529. 1888) in the lack of pruina,
distinct yellow color and flatter, shorter, more branched lobes.
Additional specimen examined. S.W.A./Namibia: Same locality as the
holotype, Nash s.n. (ASU, US).
Xanthoparmelia esterhuyseniae Hale, sp. nov. Fig. 18
Thallus laxe adnatus, saxicola vel terricola, 4-6 cm latus, lobis
sublinearibus, tenuissimis, 0.1-0.3 mm latis, ramis ultimis fere
subteretibus, superne emaculatus, isidiis sorediisque destitutus, subtus
planus, niger, sparsissime rhizinosus.
Thallus loosely adnate to free growing on soil or rocks, 4-6 cm
broad, dark yellowish green to blackening with age; lobes sublinear,
elongate, 0.1-0.3 mm wide, dichotomously branched with the ultimate lobes
nearly subterete, subascending; upper surface continuous, shiny,
emaculate; isidia and soredia absent; lower surface plane, rarely brown
or mottled at the tips but black at the center, shiny, very sparsely
rhizinate, the rhizines 0.2-0.4 mm long, black.
Pycnidia and apothecia not found.
Chemistry: Stictic, constictic, cryptostictic, norstictic and usnic
acids.
Type collection: Sandy flat rock surface on plateau, Hexburg
Mountains, elev. 1500 m, Clanwilliam Div., Cape Province, S. Africa, Grid
3319 CB, E. Esterhuysen 18476, 24 March 1951 (BOL, holotype; US,
isotype).
This finely branched species, almost looking like a soil-inhabiting
Bryoria, usually has subterete ultimate branched, similar to those of X.
molliuscula (Ach.) Hale, which is pale brown below and has broader
lobes. Another stictic acid-containing species, X. suberadicata (des
Abb.) Hale from Madagascar, has broader lobes (to 1 mm wide) and lacks
rhizines. The species is named in honor of E. Esterhuysen, who has hiked
to the tops of so many difficult high peaks in southwestern Cape Province
and discovered a number of new species in the Parmeliaceae.
Additional specimens examined S. Africa. Cape Province: Mt.
Superior, Waaihoek Mtns., Grid 3319 AD, Esterhuysen 18236 (BOL);
Platteklip near Stellenbosch, Grid 3318 DD, Garside 5037 (BOL, US).
Xanthoparmelia evernica Hale, sp. nov. Fig. 19
Thallus arcte adnatus, saxicola, 3-5 cm latus, lobis sublinearibus,
0.8-1 mm latis, superne emaculatus, aetate albo-pruinosus, isidiatus,
isidiis grosse inflatis, globosis, subtus planus, pallide brunneus,
modice rhizinosus.
Thallus tightly adnate on rock, 3-5 cm broad, darkish yellow green;
lobes sublinear, 0.8-1 mm wide, sparsely dichotomously branched, con-
tiguous, somewhat inflated; upper surface continuous, emaculate, shiny to
white pruinose, transversely cracked and rugose with age, isidiate, the
isidia coarse, globose and basally constricted, hollow, breaking
578
open at the tips, 0.2-1 mm in diameter, 0.5-1.2 mm high; lower surface
plane, pale brown to brown, moderately rhizinate, the rhizines pale
brown, 0.3-0.5 mm long.
Pycnidia and apothecia not seen.
Chemistry: Evernic, lecanoric (trace) and usnic acids.
Type collection: Laguneberg Mountains, NE of Mile 72, Distr.
Omaruru, S.W.A./Namibia, Grid 2114 CC, M. E. Hale 75106, 8 Jan. 1986 (US,
holotype; PRE, isotype).
This is another Namibian endemic clearly distinguished by the
unusually large, hollow globose isidia. The pruinose surface and
somewhat inflated aspect resemble X. serusiauxii Hale described below but
the lower surface color is different.
Additional specimens examined. S.W.A./Namibia. Same locality as
holotype, Hale 75105, 75110 (US); Hottentot site, E of Cape Cross, Grid
2114 CA, Hale 75120 (US).
Xanthoparmelia globisidiosa Hale, sp. nov. Fig. 20
Thallus adnatus, saxicola, fragilis, 4-8 cm latus, lobis subirregu-
laribus, 1-2 mm latis, superne emaculatus, isidiatus, isidiis densis,
globosis, subtus planus, pallide brunneus, rhizinis nullis.
Thallus adnate on rock, fragile and easily breaking apart when
collected, 4-8 cm broad, light yellowish green; lobes subirregular,
short, imbricate, 1-2 mm wide; upper surface continuous, shiny,
emaculate, isidiate, the isidia dense, globose, basally constricted and
breaking off apically, unbranched, 0.10-0.15 mm in diameter, 0.15-0.2 mm
high; lower surface plane, shiny, very pale tan, rhizines lacking.
Pycnidia and apothecia lacking.
Chemistry: Stictic, constictic and usnic acids, unknown substance.
Type collection: On large sandstone boulders in pasture, 25 km S of
Clanwilliam along Hwy N7, elev. ca 300 m, Cape Province, S. Africa, Grid
3218 DD, M. E. Hale 72080, 25 Jan. 1986 (US, holotype; PRE, isotype).
The most unusual features of this lichen are the lack of rhizines,
the very pale brown lower surface color and the globose isidia. It can
only be compared with X. treurensis Hale from the eastern Drakensbergs
(see below), which is coriaceous and maculate and has a black lower
surface and cylindrical isidia. This species is known only from the type
locality in Namaqualand.
Xanthoparmelia gyrophorica Hale, sp. nov. Fig. 21
Thallus adnatus, saxicola, coriaceus, 6-12 cm latus, lobis
subirregularibus, 2-6 mm latis, superne emaculatus, isidiis sorediisque
destitutus, subtus planus, pallide brunneus, modice rhizinosus.
Thallus adnate to loosely adnate on rock, firm, 6-12 cm broad,
bright yellowish green; lobes subirregular, apically rotund, contiguous
to imbricate, 2-6 mm wide; upper surface continuous, dull, emaculate,
transversely cracked with age; isidia and soredia absent; lower surface
plane, pale brown, moderately rhizinate, the rhizines brown to
blackening, rather coarse, 0.5-1 mm.
Pycnidia numerous; conidia bifusiform, 0.5 X 5-6 ym. Apothecia
common, adnate, 5-8 mm in diameter; hymenium 42-45 um; spores 6-7 X
9-10 wm.
Chemistry: Gyrophoric, lecanoric (trace), 2,4-di-O-methylgyrophoric
(trace), 5-O-methylhiascic (trace) and usnic acids (determined by C. F.
Culberson and A. Johnson).
Type collection: Exposed dolerite ledges in pasture east of Customs
579
Figures 17-20. Holotypes of Xanthoparmelia: 17, X. equalis; 18, Ke
esterhuyseniae; 19, X. evernica (lower insert X10); 20, X. globisidiosa.
Scale = mm.
580
Gate, Sani Pass, elev. 2875 m, Lesotho, Grid 2929 CB, M. E. Hale 74034,
19 Feb. 1986 (US, holotype; PRE, isotype).
Gyrophoric acid is extremely rare in Xanthoparmelia, so far known as
the main constituent only in X. leucostigma Brusse, an effigurate—
maculate, sublinear lobed species and in X. endochromatica Hale (above),
a small tightly adnate lichen with an orange-red medulla. This
conspicuous species is very common at Sani Pass and has also been found
in the Cathedral Peak area to the north. It will certainly be collected
in similar high elevation localities in the eastern escarpment of the
Drakensbergs.
Additional specimens examined. Lesotho: Same locality as the
holotype, Hale 74032, 74035, 74038, 74039, 74047 (US). S. Africa. Natal:
Cathedral Peak area, Grid 2829 CC, Schelpe s.n. (BOL).
Xanthoparmelia hybrida Hale, sp. nov. Fig. 22
Thallus adnatus, saxicola, 6-9 cm latus, lobis subirregularibus,
1-2.5 mm latis, superne emaculatus, valde transversim fissurinus, isidiis
sorediisque destitutus, subtus planus, pallide brunneus, modice
rhizinosus.
Thallus adnate on rock, 6-9 cm broad, yellowish green; lobes sub-
irregular to sublinear, 1-2.5 mm wide, contiguous to imbricate; upper
surface continuous, shiny, emaculate, becoming strongly transversely
cracked toward the center; isidia and soredia lacking; lower surface
plane, pale brown, moderately to rather densely rhizinate, the rhizines
brown, 0.5-1 mm long.
Pycnidia common; conidia bifusiform, 0.5 X 5-6 pm. Apothecia
adnate, 2-4 mm in diameter; hymenium 45-50 ym; spores not developed.
Chemistry: Protocetraric, stictic, constictic and usnic acids.
Type collection: On sandstone boulder, Treur River near Bourkes
Luck, Pilgrims Rest, elev. 1200 m, Transvaal, South Africa, Grid 2430 DB,
M. E. Hale 72083, 14 Jan. 1986 (US, holotype; PRE, isotype).
This rare lichen in the Transvaal Drakensbergs is distinguished by
the unique combination of acids, not previously reported in the genus.
Morphologically it might be mistaken for a robust specimen of X.
cumberlandia (Gyel.) Hale, an American species with stictic acid which
also occurs rarely in Cape Province.
Xanthoparmelia indumenica Hale, sp. nov. Fig. 23
Thallus arcte adnatus, saxicola, 4-5 cm latus, lobis brevi-
sublinearibus, 0.5-1.5 mm latis, superne emaculatus, isidiis sorediisque
destitutus, subtus planus, niger, sparse rhizinosus.
Thallus tightly adnate on rock, 4-5 cm broad, yellowish green but
becoming darker green toward the center; lobes sublinear, 0.5-1.5 mm
wide, rather short, subimbricate; upper surface continuous, shiny,
emaculate; isidia and soredia absent; lower surface plane, black,
moderately rhizinate, the rhizines black, rather coarse, 0.1-0.3 mm long.
Pycnidia well developed; conidia bifusiform, 0.5 X 5-6 pm.
Apothecia numerous, adnate, 0.6-1 mm in diameter; hymenium 45 pm;
spores 5-6 X 9-11 mm.
Chemistry: Stictic, constictic, cryptostictic, norstictic,
diffractaic and usnic acids (determined by C. Culberson and A. Johnson).
Type collection: On sandstone boulder, trail to Indumeni Forest/
Rainbow Gorge, Cathedral Peak, elev. 1600 m, Natal, S. Africa, Grid 2829
CC, M. E. Hale 75107 , 22 Feb. 1986 (US, holotype; PRE, isotype).
This inconspicuous lichen in the foothills of the Drakensbergs is
yep e
RES ERESSERPS ERE EER SEEDER ERR BESS
SPL PPESEER REGS
Pi 2
ae
PLAT EEE RELEEPEELAEE) Che tae
Figures 21-24. Holotypes of Xanthoparmelia: 21, X. gyrophorica; 22,
X. hybrida; 23, X. indumenica; 24, K. karooensis. Scale = mm.
582
characterized by the tightly adnate sublinear lobes, black lower surface
and the unique chemistry. Diffractaic acid had not previously been known
to occur with stictic acid in the genus. Externally it is comparable to
X. adhaerens (Nyl.) Hale, which is subcrustose with lobes less than 0.8
mm wide and contains only stictic and constictic acids.
Xanthoparmelia karooensis Hale, sp. nov. Fig. 24
Thallus arcte adnatus, saxicola, 4-7 cm latus, lobis subirregular-
ibus, brevibus, 0.8-1.5 mm latis, centrum versus congestis, bullatis,
superne emaculatus, isidiis sorediisque destitutus, subtus planus,
pallide brunneus, modice rhizinosus.
Thallus nearly subcrustose to tightly adnate on rock, 4-7 cm broad,
yellowish green; lobes subirregular, short, 0.8-1.5 mm wide, contiguous,
becoming crowded and bullate at the center; upper surface continuous,
shiny, emaculate; isidia and soredia absent; lower part of medulla ochre;
lower surface plane, pale brown, moderately rhizinate, the rhizines pale
brown, up to 0.5 mm long.
Chemistry: Hypoprotocetraric, 4-O-demethylnotatic and usnic acids,
skyrin and several unidentified minor components.
Pycnidia common; conidia bifusiform, 0.5 X 5-7 um. Apothecia num-
erous, adnate, 2-3 mm in diameter; hymenium 44-48 wm; spores 6 X 9 ym.
Type collection: Vertical schist outcrop 8 km S of Lainsburg on Hwy
R323, elev. 700 m, Cape Province, S. Africa, Grid 3320 BB, 30 Jan. 1986,
M. E. Hale 75102 (US, holotype; PRE, isotype).
This is the only tightly adnate species with hypoprotocetraric acid
and skyrin in the genus. It occurs over a wide area in Namaqualand and
southwestern Cape Province. Xanthoparmelia subdomokosii (Hale) Hale has
the same chemistry but is much larger and loosely adnate.
Additional specimens examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 75118 (US); 23 km NNW of Garies, Grid 3017
BB, Hale 72118 (US); 25 km S of Clanwilliam, Grid 3218 DD, Hale 72120
(US); Vanrhyns Pass, Grid 3119 AC, Hale 73129 (US); Anenouspas, Grid 2917
BA, Hale 73121 (US).
Xanthoparmelia lagunebergensis Hale, sp. nov. Fig. 25
Thallus arcte adnatus, saxicola, 2-4 cm latus, lobis sublinearibus,
0.5-1 mm latis, superne emaculatus, isidiis sorediisque destitutus,
subtus planus, pallide brunneus, modice rhizinosus.
Thallus tightly adnate on rock, yellowish green, 2-4 cm broad; lobes
sublinear, 0.5-1 mm wide, appearing somewhat inflated, separate to
contiguous; upper surface continuous, dull, emaculate; isidia and soredia
absent; lower surface plane, very pale brown, moderately rhizinate, the
rhizines pale, 0.2-0.4 mm long.
Pycnidia and apothecia lacking.
Chemistry: Protocetraric and usnic acids.
Type collection: Laguneberg Mountains, NE of Mile 72, Distr.
Omaruru, S.W.A./Namibia, Grid 2114 CC, M. E. Hale 75101, 8 Jan. 1986 (US,
holotype; PRE, isotype).
This Namibian endemic is close to X. serusiauxii Hale described
below but has a pale lower surface and completely different chemistry and
lacks a black rim around the lobe margins.
Xanthoparmelia lesothoensis Hale, sp. nov. Fig. 26
Thallus adnatus, saxicola, 3-5 cm latus, lobis sublinearibus,
583
LELEGE
pees
Figures 25-28, Holotypes of Xanthoparmelia: 25, X. lagunebergensis;
26, X. lesothoensis; 27, X. lividica (lower insert X10); 28, X. lobulifera.
Scale = mm.
584
brevibus, 0.8-1.5 mm latis, nigromarginatis, superne emaculatus, isidiis
sorediisque destitutus, subtus planus, brunneus, modice rhizinosus.
Thallus adnate to tightly adnate on rock, leathery, 3-5 cm broad,
dull yellowish green; lobes sublinear, 0.8-1.5 mm wide, short,
dichotomously branched, black rimmed, contiguous to subimbricate; upper
surface continuous, shiny, emaculate, transversely cracked with age,
isidia and soredia absent; lower surface plane, brown to dark brown,
moderately to densely rhizinate, the rhizines coarse, brown, 0.3-0.5 mm
long.
Pycnidia common; conidia bifusiform, 0.6 X 6-8 mm. Apothecia not
seen.
Chemistry: Salazinic, diffractaic and usnic acids (determined by C. F.
Culberson and A. Johnson).
Type collection: Exposed dolerite ledges, pasture east of the
Customs Gate, Sani Pass, elev. 2875 m, Lesotho, Grid 2929 CB, M. E. Hale
74098, 17 Feb. 1986 (US, holotype; PRE, isotype).
This small, coriaceous lichen has no close relatives. The
combination of salazinic and diffractaic acids is unusual in Xantho-
parmelia. It is obviously a high elevation species in the Drakensberg
escarpment.
Xanthoparmelia lividica Hale, sp. nov. Fig. 27
Thallus arctissime adnatus, saxicola, 2-4 cm latus, lobis sub-
linearibus, 0.1-0.2 mm latis, nigromarginatis, superne emaculatus,
isidiis sorediisque destitutus, subtus planus, niger, modice rhizinosus.
Thallus very tightly adnate on rocks, 2-4 cm broad, yellow green at
the margins but becoming darker, almost black at the center; lobes
sublinear, 0.1-0.2 mm wide, black rimmed, contiguous to crowded at the
center; surface continuous, shiny, emaculate; isidia and soredia absent;
lower surface plane, black, shiny, moderately rhizinate, the rhizines
black, ca 0.1 mm long.
Pycnidia and apothecia not found.
Chemistry: Lividic, colensoic and usnic acids (with two other faint
spots in the colensoic acid series).
Type collection: On Table Mountain sandstone ledges in Fynbos above
ocean, Olifantsbaai, Cape of Good Hope Nature Reserve, elev. 20 m, Cape
Province, S. Africa, Grid 3418 AD, M. E. Hale 74092, 21 Jan. 1986 (US,
holotype; PRE, isotype).
This tiny lichen, with perhaps the smallest lobes of any
Xanthoparmelia, is barely discernible on the surface of coarse sandstone
ledges. In size and general morphology it is closest to X. olivetorica
Hale (below), but the chemistry separates it from this and all other
species of Xanthoparmelia.
Xanthoparmelia lobulifera Hale, sp. nov. Fig. 28
Thallus arcte adnatus, saxicola, 5-8 cm latus, lobis sublinearibus,
0.6-1 mm latis, congestis, convexis, superne emaculatus, isidiis
sorediisque destitutus, subtus planus, niger, rhizinis nullis.
Thallus tightly adnate on rock, firm, 5-8 cm broad, bright yellow
green; lobes sublinear, 0.6-1 mm wide, imbricate and congested, branching
into narrow secondary lobes 0.4-0.6 mm wide, appressed to subascending;
upper surface continuous, dull to shiny, convex, emaculate, rugose and
transversely cracked with age; isidia and soredia absent; lower surface
plane to slightly grooved, black, dull, rhizines lacking.
Pycnidia and apothecia lacking.
585
Chemistry: Stictic, constictic and usnic acids.
Type collection: On sandstone in pasture, 37.4 km NW of Queenstown
along Hwy R30, elev. ca 1600 m, Cape Province, S. Africa, Grid 3126 AC,
M. E. Hale 72025, 8 Feb. 1986 (US, holotype; PRE, isotype).
There seem to be no close relatives for this eastern Cape Province
lichen. The black lower surface without rhizines and the leathery,
crowded lobes are distinctive.
Additional specimen examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 72029 (US).
Xanthoparmelia minuta Knox and Hale, sp. nov. Fig. 29
Thallus arcte adnatus, saxicola, 1-1.5 cm latus, lobis sublinear-
ibus, 0.4-0.8 mm latis, nigromarginatis, superne emaculatus, isidiatus,
isidiis cylindricis, subtus planus, niger, modice rhizinosus.
Thallus tightly adnate on rocks, 1-1.5 cm broad, yellowish green;
lobes sublinear, dichotomously branched, 0.4-0.8 mm wide, short, black
rimmed, contiguous; upper surface continuous, shiny, emaculate, isidiate,
the isidia sparse to moderate, cylindrical, black tipped, unbranched,
0.06-0.07 mm in diameter, 0.09-0.1 mm high; lower surface plane, black,
moderately rhizinate, the rhizines black, 0.3-0.5 mm long, simple.
Pycnidia and apothecia lacking.
Chemistry: Norstictic and salazinic acids in nearly equal concen-
tration, consalazinic, gyrophoric (trace), protoconstipatic (trace) and
usnic acids (determined by T. Nash, J. Elix and J. Johnston).
Type collection: Open sandstone cliffs, Mt. Sheba Nature Reserve,
Pilgrims Rest, elev. 1965 m, Transvaal, S. Africa, Grid 2480 DC, M. E.
Hale 72108, 15 Jan. 1986 (US, holotype; PRE, isotype).
This inconspicuous species, highly localized at higher elevations in
the Transvaal Drakensberg escarpment, is reminiscent of X. mougeotina
(Nyl.) Galloway, but the lobes are consistently narrower and the
chemistry is quite different (X. mougeotina has stictic acid).
Additional specimens examined. S. Africa. Transvaal: Same locality
as the holotype, Hale 72116, 72117 (US); near Houbosdorp, Magoebaskloof,
Grid 2330 CC, Nash 23337 (ASU, US). Natal: Umtanvuna Forest Reserve, Grid
3130 AA, Knox 269 (J, US).
Xanthoparmelia mollis Hale, sp. nov. Fig. 30
Thallus laxe adnatus, saxicola, mollis, usque ad 20 cm latus, lobis
lineari—elongatis, 0.5-0.7 mm latis, apice vix teretibus, superne
indistincte maculatus, isidiis sorediisque destitutus, subtus planus,
niger, rugosus, sparse rhizinosus.
Thallus loosely adnate on flat rocks, soft, forming extensive
centrifugate colonies up to 20 cm broad, yellowish green; lobes
linear-elongate, dichotomously branched, 0.5-0.7 mm wide, the ultimate
lobes nearly terete; upper surface becoming indistinctly maculate, shiny;
isidia and soredia absent; lower surface plane, black, rugose, rhizines
almost completely lacking except for a few at lobe tips.
Pycnidia and apothecia lacking.
Chemistry: Protocetraric and usnic acids.
Type collection: On flat sandstone exposures back from cliff edge,
Oribi Gorge Nature Reserve, elev. 300 m, Natal, S. Africa, Grid 3030 CB,
M. E. Hale 74048, 3 Feb. 1986 (US, holotype; PRE, isotype).
This species is in the X. hypoleia group but the thallus forms soft
centrifugate colonies, the lobes are uniformly narrow and the surface is
not strongly effigurate-maculate. It was found in abundance with X.
586
subruginosa Hale (below) on flat, exposed ledges at Oribi Gorge.
Additional specimen examined. S. Africa. Natal: Same locality as
holotype, Hale 74017 (US).
Xanthoparmelia namakwa Hale, sp. nov. Fig. 31
Thallus adnatus, saxicola, coriaceus, 4-8 cm latus, lobis sub-
linearibus, contiguis, 1-2 mm latis, superne effigurato-—maculatus,
isidiis sorediisque destitutus, subtus planus, pallide brunneus, modice
rhizinosus.
Thallus adnate on rock, rather firm and leathery, 4-8 cm broad,
darkish yellow-green; lobes sublinear, 1-2 mm wide, contiguous; upper
surface strongly effigurate-—maculate, the maculae elongate, sometimes
coalescing, shiny, becoming rugose with age toward the center; isidia and
soredia absent; lower surface plane to slightly concave, uniformly pale
brown or turning nearly black only at the tips, sparsely to moderately
rhizinate, the rhizines coarse, 0.5-1 mm long.
Pycnidia abundantly developed; conidia bifusiform, 0.5 X 5-6 mum.
Apothecia common, substipitate, 1-5 mm in diameter; hymenium 45-48 pm;
spores 5-6 X 9-11 um.
Chemistry: Salazinic and norstictic acids in nearly equal
concentration, usnic acid, with or without traces of the chalybaeizans
unknown, or rarely with salazinic acid alone.
Type collection: On schist outcrops in flat pasture land, 19.5 km
NNW of Vanrhynsdorp on east side of Hwy N7, elev. 200 m, Cape Province,
S. Africa, Grid 3118 BC, M. E. Hale 73072, 26 Jan. 1986 (US, holotype;
PRE, isotype).
This Namaqualand karoo lichen is close to X. effigurata Hale
(above), which has a uniformly black lower surface and predominantly
salazinic acid and the chalybaeizans unknown. The two species have only
been collected together at Anenouspas. Another closely related species,
X. exornata (Stein. & Zahlbr.) Knox, has strongly convex lobes, coarse
rhizines and recessed, pseudocyphella-like maculae.
Additional specimens examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 73061, 73071 (US); Anenouspas, Grid 2917
BA, Hale 73062, 73070 (US); 44 km N of Vanrhynsdorp, Grid 3118 BC, Hale
73073 (US); 4.9 km S of Lainsburg, Grid 3320 BB, Hale 73074 (US).
Xanthoparmelia namaquensis Hale, sp. nov. Fig. 32
Thallus laxe adnatus, saxicola, 6-12 cm latus, lobis sublinearibus,
2-5 mm latis, superne continuus vel vix maculatus, isidiis sorediisque
destitutus, medulla partim purpurea, subtus planus, niger, sparse
rhizinosus.
Thallus loosely adnate on rock, easily breaking apart, 6-12 cm
broad, yellowish green but darkening at the center; lobes sublinear,
rather short and crowded, 2-5 mm wide, contiguous to imbricate; upper
surface continuous to very weakly maculate, shiny, transversely cracked
with age; isidia and soredia absent; lower part of medulla in part dark
purple; lower surface plane, black, dull, sparsely rhizinate, the
rhizines black, coarse, 0.5-1 mm long.
Pycnidia and apothecia lacking.
Chemistry: Lecanoric and usnic acids, a bright pinkish orange
anthraquinone pigment on extraction with acetone (detected only in
decomposing parts of the medulla) (determined by C. F. Culberson and A.
Johnson).
Type collection: Exposed sandstone cliff on edge of escarpment, 41
587
a,
Figures 29-32. Holotypes of Xanthoparmelia: 29, X. minuta; 30, X.
mollis; 31, X. namakwa (lower insert X10); 32, X. namaquensis. Scale = mm.
588
km NE of Vanrhynsdorp at Vanrhyns Pass along Hwy R27, elev. 800 m,Cape
Province, S. Africa, Grid 3119 AC, M. E. Hale 73018, 28 Jan. 1986 (US,
holotype; PRE, isotype). é
Of the eight species of Xanthoparmelia with lecanoric acid, this is
the only one that is clearly loosely adnate. It produces a unique deep
pinkish orange pigment reacting K+ light purple and probably related (but
not equal) to the anthraquinone series in X. endomiltoides. It is
apparently restricted to Namaqualand.
Additional specimens examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 73131 (US); 1 km S of Nuwerus, Grid 3118
AB, Hale 72114 (US).
Xanthoparmelia neosynestia Hale, sp. nov. Fig. 33
Thallus adnatus vel laxe adnatus, saxicola, 4-7 cm latus, lobis
sublinearibus, 1-2.5 mm latis, superne maculatus, isidiis sorediisque
destitutus, subtus planus, pallide brunneus, modice rhizinosus.
Thallus adnate to loosely adnate on rock, rather brittle, dark
greenish yellow, 4-7 cm broad; lobes sublinear to subirregular, 1-2.5 mm
wide, short and imbricate; upper surface uniformly maculate, shiny,
transversely cracked with age; isidia and soredia absent; lower surface
plane, pale brown to brown, sparsely to moderately rhizinate, the
rhizines brown, 0.5-1 mm long, simple.
Pycnidia numerous; conidia bifusiform, 0.56 X 5-6 ym. Apothecia well
developed, adnate, 3-5 mm in diameter, the rim crenate; hymenium 40-45
yum; spores 5-6 X 9-10 um.
Chemistry: Salazinic, norstictic (trace), protocetraric (trace),
consalazinic and usnic acids, with or without the chalybaeizans unknown
(present in holotype) (determined by T. Nash and J. Elix).
Type collection: Exposed sandstone cliffs on escarpment in hillside
karoo, 41 km NE of Vanrhynsdorp at Vanrhyns Pass on Hwy R27, elev. 800
m, Cape Province, S. Africa, Grid 3119 AC, M. E. Hale 73017, 28 Jan. 1986
(US, holotype; PRE, isotype).
This species has the same distinctive dull yellowish green cast of
X. synestia (Stirt.) Hale but the lower surface is uniformly pale or
darker brown but not blackening. It occurs over a broad area in
Namaqualand and southwestern Cape Province.
Additional specimens examined. S. Africa. Cape Province: 1 km E of
Springbok, Grid 2917 DB, Hale 72095 (US); Pakhuis Pass, Grid 3218 BB,
Hale 72119 (US); Camps Bay, Grid 3318 CD, Almborn 4393, 4395 (LD, US);
Bainskloof Pass, Grid 3319 CA, Hale 72121 (US), Nash 23519 (ASU, US),
Sipman 20234 (B); 26.4 km E of Ladismith, Grid 3321 AD, Hale 73144 (US);
Meiringspoort, Grid 3322 BC, Hale 73143 (US).
Xanthoparmelia neotasmanica Hale, sp. nov. Fig. 34
Thallus laxe adnatus, saxicola, 10-15 cm latus, lobis sublinear-
ibus, 2-2.5 mm latis, centrum versus laciniatis, superne leviter
maculatus, isidiis sorediisque destitutus, subtus planus, niger, modice
rhizinosus.
Thallus loosely adnate, often growing in mats on tops and sides of
sheltered rocks, 10-15 cm broad, light yellowish green; lobes sublinear,
2-2.5 mm wide, branching into numerous short laciniae about 0.6-1.0 mm
wide toward the center; upper surface weakly to distinctly maculate,
shiny, isidia and soredia absent; lower surface plane, black, moderately
to densely rhizinate, the rhizines black, 0.5-1 mm long.
Pycnidia abundant; conidia bifusiform, 0.6 X 6-7 am. Apothecia
589
a
rr a3he fas
Figures 33-36. Holotypes of Xanthoparmelia: 33, X. neosynestia; 34,
X. neotasmanica; 35, X. nuwarensis; 36, X. ochropulchra. Scale = mm.
590
rather rare, adnate, 4-9 mm in diameter; hymenium 45-48 pm; spores 5-6
X 9-10 mm (data from Hale 73019).
Chemistry: Salazinic and usnic acids.
Type collection: Exfoliating granitic outcrops in karoo, 78.6 km S
of Sutherland on Hwy R354, Cape Province, S. Africa, Grid 3220 DC, M. E.
Hale 73019, 29 Jan. 1986 (US, holotype; PRE, isotype).
The X. tasmanica group encompasses a wide range of variation in lobe
width and surface features. Xanthoparmelia tasmanica (Hook. & Tayl.)
Hale itself is a very common species in S. Africa, as well as in North
America and Australasia. This new species represents one extreme,
consistently producing marginal laciniae and having fairly distinct,
uniform maculae, characters lacking in typical X. tasmanica. It has a
wide range in southwestern Cape Province.
Additional specimens examined. S. Africa. Cape Province: Lions Head,
Cape Div., Grid 3318 AB, Almborn & Karnefelt 8411-10 (LD); Moltenopas,
Grid 3222 BA, Hale 73006 (US); Nieuwoudtville Nature Reserve, Grid 3119
AC, Hale 72012 (US); Meiringspoort, Grid 3322 BC, Hale 73005 (US).
Xanthoparmelia nuwarensis Hale, sp. nov. Fig. 35
Thallus laxe adnatus, saxicola, 6-9 cm latus, lobis sublinearibus,
1.5-3 mm latis, apice subadscendentibus, superne valde maculatus, subtus
planus, brunneus, sparse rhizinosus.
Thallus loosely adnate on rocks, breaking apart, 6-9 cm broad, dark
greenish yellow; lobes sublinear, 1.5-3 mm wide, dichotomously branched,
separate to imbricate, the tips curling up somewhat; upper surface
strongly maculate, shiny; isidia and soredia absent; lower surface plane,
becoming rugose in part, black at the tips but brown to pale brown toward
the center, sparsely to moderately rhizinate, the rhizines brown, 0.5-1
mm long, rather coarse.
Pycnidia commonly developed; conidia bifusiform, 0.5 X 6-7 mm.
Apothecia substipitate, 3-6 mm in diameter; hymenium 43-46 jm; spores
5-6 X 9-11 um.
Chemistry: Fumarprotocetraric, succinprotocetraric and usnic acids.
Type collection: Large exposed sandstone ledge and ridges in
pasture, 1 km S of Nuwerus on Hwy 368, elev. ca 500 m, Cape Province, S.
Africa, Grid 3118 AB, M. E. Hale 72111, 26 Jan. 1986 (US, holotype; PRE,
isotype).
This Namaqualand lichen is a member of the highly variable X.
phaeophana (Stirt.) Hale complex. It is differentiated by the darker
green color, strong white maculation as in X. synestia (Stirt.) Hale, and
the rugose, brown lower surface. Xanthoparmelia viridis Hale (below) is
very close and has the same chemistry but differs in being black below.
Additional specimen examined. S. Africa. Cape Province: 23 km NNW of
Garies, Grid 3017 BB, Hale 72112 (US).
Xanthoparmelia ochropulchra Hale, sp. nov. Fig. 36
Thallus arcte adnatus, saxicola, friabilis, 3-8 cm latus, lobis
subirregularibus, 0.6-1.4 mm latis, superne emaculatus, isidiis
sorediisque destitutus, medulla pro parte ochracea, subtus planus,
pallide brunneus, sparse rhizinosus.
Thallus tightly adnate, soft and friable, 3-8 cm broad, yellowish
green; lobes subirregular, 0.6-1.4 mm wide, contiguous to subimbricate;
upper surface continuous, shiny, weakly reticulate-maculate and minutely
rugulose at the tips; isidia and soredia absent; medulla pale orange in
the lower parts; lower surface plane, pale brown, sparsely rhizinate, the
So
the rhizines pale brown, 0.2-0.4 mm long.
Pycnidia common; conidia bifusiform, 0.6 X 5-6 mm. Apothecia well
developed, adnate, 1-1.5 mm in diameter; hymenium 40-43 um; spores 5 X
8-9 um.
Chemistry: A new depside related to diffractaic acid and usnic acid,
endocrocin and an unidentified anthraquinone pigment (determined by C. F.
Culberson and A. Johnson).
Type collection: Exfoliating granitic outcrops in karoo, 78.6 km S
of Sutherland on Hwy R354, elev. ca 900 m, Cape Province, S. Africa, Grid
3220 DC, M. E. Hale 73101, 29 Jan. 1986 (US, holotype; PRE, isotype).
Known only from one locality in the karoo of southwestern Cape
Province, this species has no obvious relatives in Xanthoparmelia. The
yellow pigments, presence of a new depside near diffractaic acid (also
present in X. duplicata Hale (above)) and small, friable, short-lobed
thallus are all unique.
Xanthoparmelia olifantensis Hale, sp. nov. Fig. 37
Thallus arcte adnatus, 2-3 cm latus, lobis sublinearibus, 0.4-0.6 mm
latis, centrum versus subareolatis, superne emaculatus, isidiis sorediis—
que destitutus, subtus planus, niger, sparse rhizinosus.
Thallus tightly adnate to almost subcrustose, 2-3 cm broad, yellow-
ish green; lobes sublinear, 0.4-0.6 mm wide, sparsely dichotomously
branched, cracked and subareolate toward the center; upper surface
continuous, shiny, emaculate, transversely cracked; isidia and soredia
absent; lower surface plane, black, sparsely rhizinate, the rhizines
black, 0.2-0.3 mm long.
Pycnidia and apothecia lacking.
Chemistry: Norstictic, salazinic and usnic acids.
Type collection: On ledges above beach, Olifantsbaai, Cape of Good
Hope Nature Reserve, elev. 20 m, Cape Province, S. Africa, Grid 3418 AD,
M. E. Hale 72113, 21 Jan. 1986 (US, holotype; PRE, isotype).
A similar species is X. minuta Knox & Hale (see above), which also
contains norstictic and salazinic acids in nearly equal concentration but
differs in having isidia and a trace of gyrophoric acid. This species is
apparently restricted to the rocky hillsides exposed to southwesterly
rain and mist in the Cape area.
Additional specimens examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 75119 (US); Kogelberg Range, Grid 3418 BD,
Leighton 716 (BOL).
Xanthoparmelia olivetorica Hale, sp. nov. Fig. 38
Thallus arcte adnatus, 1-3 cm latus, saxicola, lobis sublinearibus,
0.2-0.6 mm latis, valde nigromarginatis, superne emaculatus, isidiis
sorediisque destitutus, subtus planus, niger, sparse rhizinosus.
Thallus tightly adnate on rock, 1-3 cm broad, dark greenish yellow
but blackening with age toward the center; lobes sublinear, 0.2-0.6 mm
wide, strongly black rimmed, contiguous; upper surface continuous, shiny,
transversely cracked with age, isidia and soredia absent; lower surface
plane, black, shiny, sparsely rhizinate, the rhizines black, about 0.2 mm
long.
Pycnidia and apothecia lacking.
Chemistry: Olivetoric and usnic acids.
Type collection: On sheltered sandstone ledges along trail from
Platteklip Gorge to Woodhead Reservoir, Table Mountain Nature Reserve,
Cape Province, S. Africa, Grid 3318 CD, M. E. Hale 72079, 23 Jan. 1986
592
(US, holotype; PRE, isotype).
The appressed thallus with very narrow lobes resembles X. lividica
Hale (above) but is more robust. Another species with olivetoric acid,
X. heterodoxa Hale (described from Natal Table Mountain), is larger, less
tightly adnate and lacks a black margin around the lobes. These are the
only species in Xanthoparmelia with olivetoric acid.
Additional specimen examined. Same locality as the holotype, Hale
72080 (US).
Xanthoparmelia oribensis Hale, sp. nov. Fig. 39
Thallus adnatus, saxicola, 6-12 cm latus, lobis subirregularibus,
2-4 mm latis, superne emaculatus, isidiatus, isidiis densis, cylindricis,
medulla pro parte ochracea, subtus planus, niger, modice rhizinosus.
Thallus adnate to closely adnate on flat rocks, 6-12 cm broad,
yellowish green; lobes subirregular to sublinear, 2-4 mm wide, contig-—
uous; upper surface continuous, shiny, emaculate, transversely cracked
with age, isidiate, the isidia moderate to dense, tall, cylindrical,
simple or branched, 0.1-0.12 mm in diameter, 0.5-1 mm high; lower part of
medulla orange-red; lower surface plane, black but ochre when lower
cortex breaks open, sparsely to moderately rhizinate, the rhizines black,
simple, 0.5-1 mm long.
Pycnidia and apothecia lacking.
Chemistry: Hypoprotocetraric, 4-O-demethylnotatic and usnic acids,
skyrin and several minor unidentified components.
Type collection: Flat sandstone exposures back from the cliff, Oribi
Gorge Nature Reserve, elev. 300 m, Natal, S. Africa, Grid 3030 CB, M. E.
Hale 74041, 3 Feb. 1986 (US, holotype; PRE, isotype).
The only other species in Xanthoparmelia with hypoprotocetraric acid
and a black lower surface is subcrustose X. neocongensis (Hale) Hale,
which is also isidiate but lacks skyrin. Two other African species with
hypoprotocetraric acid and skyrin, X. karooensis Hale (above) and X.
subdomokosii (Hale) Hale, lack isidia. Xanthoparmelia oribensis is known
only from the spectacular sandstone cliffs overlooking Oribi Gorge.
Additional specimens examined. S. Africa. Natal: Same locality as
the holotype, Hale 74054, 74055 (US).
Xanthoparmelia probarbellata Hale, sp. nov. Fig. 40
Thallus adnatus, saxicola, 4-6 cm latus, lobis sublinearibus, 2-3 mm
latis, superne emaculatus, isidiis sorediisque destitutus, medulla pro
parte ochracea, subtus planus, pallide brunneus, modice rhizinosus.
Thallus adnate on rock, 4-6 cm broad, yellowish green; lobes
sublinear, 2-3 mm wide, somewhat separate to contiguous; upper surface
continuous, shiny, emaculate; isidia and soredia absent; medulla ochre in
the lower part; lower surface plane, pale brown to brown, moderately
rhizinate, the rhizines brown, 0.5-1 mm long.
Pycnidia numerous; conidia bifusiform, 0.56 X 5-6 um. Apothecia
numerous, adnate, 3-5 mm in diameter; hymenium 40-45 pm; spores 5-6 X
8-9 yim.
Chemistry: Salazinic and norstictic acids in nearly equal
concentration, trace of chalybaeizans unknown, usnic acid, and skyrin.
Type collection: Soft conglomerate sandstone ridges in pasture,
52.5 km SE of Lady Grey on Hwy R58, elev. ca 1900 m, Cape Province, S.
Africa, Grid 3027 CD, M. E. Hale 72109, 9 Feb. 1986 (US, holotype; PRE,
isotype).
An Australian species, X. barbellata (Kurok.) Hale, has similar
593
a
‘
Figures 37-40. Holotypes of Xanthoparmelia: 37, X. olifantensis; 38,
X. olivetorica; 39, X. oribensis; 40, X. probarbellata. Scale = mm.
594
chemistry (except for the lack of the chalybaeizans unknown) but is more
loosely adnate and may occur on soil as well as rocks. Xanthoparmelia
probarbellata is known only from the type collection in eastern Cape
Province.
Xanthoparmelia pustulifera Hale, Nash and Elix, sp. nov. Fig. 41
Thallus adnatus, saxicola, 5-8 cm latus, lobis sublinearibus,
brevibus, 0.5-1.2 mm latis, imbricatis, superne emaculatus, isidiatus,
isidiis globosis, eruptentibus pustulescentibusque, subtus planus, niger,
modice rhizinosus.
Thallus adnate to tightly adnate on rock, fragile, 5-8 cm broad,
dull yellowish green; lobes short and sublinear, dichotomously branched,
0.5-1.2 mm wide, contiguous to subimbricate, becoming sublobulate with
age, the lobules appressed; upper surface continuous, shiny, emaculate,
irregularly cracked with age, moderately isidiate, the isidia initially
globose, 0.1-0.3 mm in diameter and up to 0.3 mm high, soon erupting into
pustulate or coarsely subsorediate masses; lower surface plane, brownish
at the tips but black toward the center, moderately rhizinate, the
rhizines black, 0.2-0.3 mm long.
Pycnidia and apothecia not found.
Chemistry: Stictic, constictic, norstictic (trace) and usnic acids
and an unidentified fatty acid.
Type collection: Open dolerite ridges in karoo, 22 km ENE of Prince
Albert on the road to Seekoeigat, elev. 1000 m, Cape Province, S. Africa,
Grid 3322 AB, M. E. Hale 75115, 31 Jan. 1986 (US, holotype; PRE,
isotype).
The only closely related species, X. mougeotii (Schaer.) Hale, has
distinct capitate soralia and is more tightly adnate. I have not had an
opportunity to revise herbarium records of X. mougeotii from S. Africa;
these may well represent X. pustulifera.
Additional specimen examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 75116 (US); Devil's Peak, Table Mountain,
Grid 3318 CD, Nash s.n. (ASU).
Xanthoparmelia rubromedulla Hale, sp. nov. Fig. 42
Thallus laxe adnatus, saxicola, 5-10 cm latus, lobis irregulariter
sublinearibus, 1-1.5 mm latis, superne distincte maculatus, isidiis
sorediisque destitutus, subtus planus, niger, modice rhizinosus.
Thallus loosely adnate on rock, rather rigid and brittle, frag-
menting, 5-10 cm broad, light yellowish green; lobes sublinear, 1-1.5 mm
wide, dichotomously branched, subimbricate; upper surface uniformly
maculate, shiny, isidia and soredia absent; lower part of medulla
orange-red; lower surface plane, brown at the tips but becoming black,
sparsely to moderately rhizinate, the rhizines black, rather coarse,
0.5-1 mm long.
Pycnidia sparsely developed; conidia not found. Apothecia absent.
Chemistry: Salazinic and usnic acids, skyrin.
Type collection: Sandy rock face above stream, Waaihoek Peak, elev.
1680 m, Worcester Division, Cape Province, S. Africa, Grid 3319 AD, E.
Esterhuysen 22560, 12 Jan. 1954 (BOL, holotype; US, isotype).
This species has some resemblance to X. synestia (Stirt.) Hale but
the lobes are more elongate, the thallus is more rigid, quite loosely
adnate and pulvinate on rock, and skyrin is present in the medulla. Both
X. luminosa (Elix) Hale and X. surrogata Hale (below) are more closely
adnate and coriaceous with broader lobes.
595
Figures 41-44, Holotypes of Xanthoparmelia: 41, X. pustulifera; 42,
X. rubromedulla; 43, X. saniensis; 44, X. serusiauxii. Scale = mm.
596
Xanthoparmelia saniensis Hale, sp. nov. Fig. 43
Thallus arcte adnatus vel fere subcrustosus, saxicola, 4-6 cm latus,
lobis subirregularibus, 1-1.5 mm_ latis, centrum versus bullatis, superne
emaculatus, isidiatus, isidiis globosis, fragilibus, subtus planus,
pallide brunneus, modice rhizinosus.
Thallus tightly adnate to nearly subcrustose on doleritic rock, 4-6
em broad, yellowish green; lobes subirregular, 1-1.5 mm wide, brown to
black rimmed, contiguous to imbricate and bullate at the center; upper
surface continuous, shiny, emaculate, isidiate, isidia dense, globose and
basally constricted, bursting open apically, 0.1-0.3 mm in diameter,
0.1-0.3 mm high; lower part of medulla ochre; lower surface plane, pale
brown tinged with ochre, moderately rhizinate, the rhizines pale brown,
0.3-0.5 mm long.
Pycnidia and apothecia lacking.
Chemistry: Hypostictic, hyposalazinic and usnic acids, another
"quintaria" unknown, and skyrin.
Type collection: Exposed dolerite ledges, Sani Pass east of Customs
Gate, elev. 2875 m, Lesotho, Grid 2929 CB, M. E. Hale 74031, 19 Feb. 1986
(US, holotype; PRE, isotype).
This remarkable high elevation Drakensberg species, characterized by
the globose isidia and tightly adnate habit, has only one probable
relative, the rare Brazilian X. kalbii Hale, another hypostictic
acid-containing Xanthoparmelia, not as tightly adnate, with broader lobes
and without skyrin.
Xanthoparmelia serusiauxii Hale, sp. nov. Fig. 44
Thallus adnatus, saxicola, 5-10 cm latus, lobis sublinearibus,
inflatis, 1-2 mm latis, superne continuus, nitidus vel pruinosus, isidiis
sorediisque destitutus, subtus planus, niger, sparse rhizinosus.
Thallus adnate to tightly adnate on rock, soft, 5-10 cm broad, dull
greenish yellow and blackening at the center with age; lobes sublinear,
little branched, 1-2 mm wide, appearing inflated, black-rimmed, con-
tiguous, short laciniate at the center with age; upper surface contin-
uous, the tips faintly reticulate—maculate, shiny or mostly dull white
pruinose, convex and heavily rugose at the center; isidia and soredia
absent; lower surface plane, black, dull, very sparsely rhizinate, the
rhizines coarse, 0.3-0.5 mm long.
Pycnidia common; conidia not found. Apothecia lacking.
Chemistry: Lecanoric and usnic acids.
Type collection: Laguneberg Mountains, NE of Mile 72, Distr.
Omaruru, S.W.A./Namibia, Grid 2114 CC, M. E. Hale 75102, 8 Jan. 1986 (US,
holotype; PRE, isotype).
This is a common Xanthoparmelia in the central Namib area. It
grows on large dolerite boulders and small flat pebbles in desert pave-
ment. The lobes are sublinear, little branched, and seem inflated. The
medulla is especially thick and soft. I examined the surface for epi-
cortical pores with the scanning electron microscope. They are present
but obscured by crystals of pruina except at lobe margins. The species
is named in honor of Dr. Emanuel Serusiaux, the first lichenologist to
collect this unusual lichen in Namibia. Related species in this unusual
locality include X. evernica Hale and X. lagunebergensis Hale (above).
Additional specimens examined. S.W.A./Namibia: Same locality as the
holotype, Hale 75108, 75109, 75111 (US), Nash 23303 (ASU, US); gravel
flats east of Cape Cross, Grid 2114 CA, Hale 751138, 75114 (US).
290
Xanthoparmelia simulans Hale, sp. nov. Fig. 45
Thallus laxe adnatus, saxicola vel terricola, 5-8 cm latus, lobis
linearibus, 0.5-1 mm latus, nigro-marginatis, irregulariter constrictis,
superne emaculatus, isidiis sorediisque destitutus, subtus planus, niger,
sparse rhizinosus.
Thallus loosely adnate on soil, 5-8 cm broad, bright yellowish
green; lobes linear, 0.5-1 mm wide, black rimmed, little branched and
irregularly constricted, divaricately branched, subascending; upper
surface continuous, shiny, emaculate; isidia and soredia absent; lower
surface plane, black, shiny, sparsely rhizinate, the rhizines black,
rather coarse, 0.5-1 mm long.
Pycnidia and apothecia lacking.
Chemistry: Stictic, constictic, cryptostictic, norstictic (trace)
and usnic acids.
Type collection: Windswept ridge on S side of Swartberg Pass,
Swartberg Mountains, NW of Kango Caves on Hwy R828, elev. 1800 m, Cape
Province, S. Africa, Grid 3322 AC, M. E. Hale 74045, 31 Jan. 1986 (US,
holotype; PRE, isotype).
This species differs from closely related X. constrictans (Nyl.)
Hale in presence of the stictic acid group (in place of salazinic acid).
While X. suberadicata (des Abb.) Hale from Madagascar also has stictic
acid, it lacks rhizines and has uniformly linear lobes with a strongly
rugose lower surface. Xanthoparmelia simulans occurs at higher
elevations in southwestern Cape Province.
Additional specimens examined. S. Africa. Cape Province: Same
locality as the holotype, Stirton 9535 (PRE); Platteklip Gorge to
Woodhead Reservoir, Grid 3318 CD, Degelius 1A-297 (UPS, US), Hale 72032,
72035, 72045 (US); 32.7 km ESE of Citrusdal, Grid 3219 CA, Hale 72028
(US).
Xanthoparmelia skyrinifera Hale, sp. nov. Fig. 46
Thallus laxe adnatus, coriaceus, 6-12 cm latus, lobis sublinear-
ibus, 1.5-2.5 mm latis, superne continuus vel sparse maculatus, isidiis
sorediisque destitutus, medulla pro parte ochracea, subtus planus,
pallide brunneus, modice rhizinosus.
Thallus loosely adnate on rocks, firm, 6-12 cm broad, bright
yellowish green; lobes sublinear, 1.5-2.5 mm wide, more or less imbri-
cate; upper surface faintly to distinctly maculate, rugose and trans-
versely cracked with age; isidia and soredia absent; medulla ochre in the
lower part; lower surface plane, pale brown with an ochre tinge but
turning darker brown toward the center, sparsely to moderately rhizinate,
the rhizines pale brown, 0.5-1.5 mm long, simple to furcate and apically
splayed.
Pycnidia (from Hale 74094) common; conidia bifusiform, 0.5 X 5-6
pm. Apothecia lacking.
Chemistry: Protocetraric and usnic acids, skyrin.
Type collection: Low schist ridge in pasture, 15.9 km NE of
Barrydale along Hwy R62, elev. ca 500 m, Cape Province, S. Africa, Grid
8321 DD, M. E. Hale 74040, 2 Feb. 1986 (US, holotype; PRE, isotype).
This unique lichen from southern Cape Province is characterized by
the loosely adnate habit, pale brown, moderately rhizinate lower surface
and chemistry. Another species with protocetraric acid and skyrin, X.
subcolorata Hale (below), has a maculate surface, rugose eroding lower
surface and sparse rhizines.
Additional specimens examined. S. Africa. Cape Province: Same
598
locality as the holotype, Hale 74102, 74105 (US); 23 km SW of Ladismith,
Grid 3321 CA, Hale 74094, 74095 (US).
Xanthoparmelia stenosporonica Hale, sp. nov. Fig. 47
Thallus arcte adnatus, saxicola, 1-2 cm latus, lobis sublinearibus,
0.2-0.5 mm latis, brevibus, nigromarginatis, superne emaculatus, isidiis
sorediisque destitutus, subtus planus, niger, modice rhizinosus.
Thallus tightly adnate on rock, 1-2 cm broad, light yellowish green;
lobes sublinear to subirregular, 0.2-0.5 mm wide, short, contiguous to
imbricate, black rimmed; upper surface continuous, shiny, emaculate;
isidia and soredia absent; lower surface plane, shiny, black, moderately
rhizinate, the rhizines black, 0.1-0.2 mm long.
Pycnidia and apothecia lacking.
Chemistry: Stenosporonic, colensoic and usnic acids.
Type collection: Summit of Swartberg Pass on Hwy R828, elev. 1585 m,
Cape Province, S. Africa, Grid 3322 AC, M. E. Hale 72117, 31 Jan. 1986
(US, holotype; PRE, isotype).
This inconspicuous species grows intermixed with X. endochromatica
Hale (above) in an exposed, high elevation habitat in southwestern Cape
Province. Stenosporonic acid has not previously been reported in the
genus.
Additional specimen examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 72082 (US).
Xanthoparmelia subcolorata Hale, sp. nov. Fig. 48
Thallus adnatus vel laxe adnatus, saxicola, 6-8 cm latus, lobis
sublinearibus, subadscendentibus, 2-4 mm latis, Superne maculatus,
isidiis sorediisque destitutus, medulla pro parte ochracea, subtus
planus, rugosus, pallide brunneus, sparsissime rhizinosus.
Thallus adnate to loosely attached on rock, forming discrete, nearly
umbilicate colonies, 6-8 cm broad, dark yellowish green; lobes sublinear,
2-4 mm wide, crowded and imbricate, subascending; upper surface
moderately to strongly maculate, shiny, deeply transversely cracked;
isidia and soredia absent; lower part of medulla ochre; lower surface
plane, pale or darker brown with ochre spots, blackening toward the tips,
strongly rugose and in part eroding with age, rhizines absent or very
sparse, brown, coarse, to 1 mm long.
Pycnidia common; conidia bifusiform, 0.56 X 5-6 um. Apothecia
common, adnate, 5-10 mm in diameter; hymenium 40-45 um; spores 5-6 X
10-12 yum.
Chemistry: Protocetraric and usnic acids, skyrin.
Type collection: Sloping sandstone boulders in Fynbos, 19.9 km NW of
Kango Caves on Swartberg Pass Road, elev. ca 1100 m, Cape Province, S.
Africa, Grid 3322 AC, M. E. Hale 73014, 31 Jan. 1986 (US, holotype; PRE,
isotype).
This loosely adnate, almost umbilicate lichen resembles X. denudata
Hale (see above), which has a similar lower surface and skyrin but
differs in having salazinic acid as the main component and narrower
lobes. It also falls close to X. austroafricana (Stirt.) Hale, which has
a more typically foliose thallus and lacks skyrin and distinct maculae.
It has a typical Cape-Namaqualand distribution with a disjunct record in
Transvaal.
Additional specimens examined. Cape Province: 1 km S of Nuwerus on
Hwy 363, Grid 3118 AB, Hale, 72107 (US); 44 km N of Vanrhynsdorp, Grid
3118 BC, Hale 73115, 75116, 75117 (US); 32.7 km ESE of Citrusdal, Grid
5999
Figures 45-48. Holotypes of Xanthoparmelia: 45, X. simulans; 46, Ks
skyrinifera; 47, X. stenosporonica (enlarged X10); 48, X. subcolorata.
Scale = mm.
600
3219 CA, Hale 72088 (US); summit of Bainskloof on Hwy Nl, Grid 3319 CA,
Hale 72104, 72106 (US), Sipman 20229 (B); Anenouspas, Grid 2917 BA, Hale
73131 (US); 26.4 km E of Ladismith, Grid 3321 AD, Hale 73057 (US);
Kruisrivier, Grid 3321 BB, Hale 73056 (US). Transvaal: Lejuma Summit,
Soutpansberg Mountains, Grid 2329 AB, Sipman 19871 (B).
Xanthoparmelia subcrustosa Hale, sp. nov. Fig. 49
Thallus arcte adnatus, subcrustosus, saxicola, 2-3 cm latus, lobis
subirregularibus, 0.6-1.0 mm latis, centrum versus areolato-—bullatis,
superne emaculatus, isidiis sorediisque destitutus, subtus planus,
pallide brunneus, modice rhizinosus.
Thallus subcrustose to tightly adnate on rocks, 2-3 cm broad, pale
yellowish green; lobes subirregular, 0.6-1.0 mm wide, bullate-areolate in
the center; upper surface continuous, shiny or at times pruinose,
emaculate, transversely cracked; isidia and soredia absent; lower surface
plane, pale brown, moderately rhizinate, the rhizines pale brown, 0.3-0.5
mm long.
Pycnidia common; conidia bifusiform, 0.6 X 5-6 mm. Apothecia poorly
developed, adnate, 1 mm in diameter; hymenium 42-45 ym; spores 6 X 8-9
ym.
Chemistry: Salazinic, norstictic, connorstictic and usnic acids,
chalybaeizans unknown.
Type collection: On granite boulders in Karoo, 8.3 km E of Garies on
Studerspas Road, elev. 900 m, Cape Province, S. Africa, Grid 3018 AC, M.
E. Hale 73016, 26 Jan. 1986 (US, holotype; PRE, isotype).
Xanthoparmelia subcrustosa is unique among subcrustose species in
chemistry. It is widely distributed in southern Cape Province.
Additional specimens examined. S. Africa. Cape Province: Ceres
Nature Reserve SW of Ceres, Grid 3319 AD, Hale 72110 (US); 78.6 km S of
Sutherland, Grid 3220 DC, Hale 73122 (US); 33.8 km W of Dordrecht, Grid
3126 BC, Hale 73119, 73120 (US).
Xanthoparmelia subnigra Hale, sp. nov. Fig. 50
Thallus adnatus vel arcte adnatus, 4-6 cm latus, lobis sublinear-
ibus, 1-2 mm latis, superne emaculatus, isidiis sorediisque destitutus,
subtus planus, niger, modice rhizinosus.
Thallus adnate to tightly adnate on rocks, 4-6 cm broad, light
yellowish green; lobes sublinear to subirregular, 1-2 mm wide, contig-
uous; upper surface continuous, shiny, emaculate; isidia and soredia
absent; lower surface plane, black, sparsely to moderately rhizinate, the
rhizines black, 0.5-1 mm long.
Pycnidia common; conidia bifusiform, 0.6 X 6-7 mm. Apothecia
numerous, adnate, 1-2 mm in diameter; hymenium 44-48 wm; spores 6-7 X
8-10 mm.
Chemistry: Dehydroconstipatic (major), pertusaric (major),
protoconstipatic (trace), constipatic (trace) and usnic acid in addition
to other unidentified fatty acids (determined by J. Johnston).
Type collection: Small sandstone outcrops in pasture, summit of Long
Tom Pass on S side of Hwy R37, Farm De Kuiten 205, elev. 2140 m,
Lydenburg, Transvaal, S. Africa, Grid 2430 BA, M. E. Hale 72026, 16 Jan.
1986 (US, holotype; PRE, isotype).
No other constipatic acid-containing species of Xanthoparmelia have
a black lower surface, excepting X. aliphatica Hale (above), which is a
much larger, loosely adnate lichen. A comparable species with a pale
lower surface is the widespread X. subdecipiens (Vain.) Hale. A
601
ie Ns
iy
Ea
Figures 49-52. Holotypes of Xanthoparmelia: 49, X. subcrustosa; 50,
X. subnigra; 51, X. subochracea; 52, X. subruginosa. Scale = mm.
602
presumptive isidiate morphotype, X. transvaalensis Hale, Nash & Elix
(below), occurs in the same locality.
Additional specimen examined. S. Africa. Transvaal: Same locality as
the holotype, Nash 234388 (ASU, US).
Xanthoparmelia subochracea Hale, sp. nov. Fig. 51
Thallus adnatus, saxicola, 3-6 cm latus, lobis subirregularibus,
1.5-2.5 mm latis, superne emaculatus, convexus, isidiis sorediisque
destitutus, medulla pro parte ochracea, subtus planus, brunneus, modice
rhizinosus.
Thallus rather closely adnate on rock, 3-6 cm broad, bright
yellowish green; lobes subirregular, 1.5-2.5 mm wide, apically rotund,
black rimmed, contiguous; upper surface continuous, shiny, emaculate,
convex, bullate toward the center; isidia and soredia absent; medulla
ochre in the lower part; lower surface plane, brown to dark brown,
sparsely to moderately rhizinate, the rhizines brown, coarse, ca 0.5 mm
long.
Pycnidia common; conidia bifusiform, 5-6 um. Apothecia poorly
developed, adnate, 1-2.5 mm in diameter; hymenium ca 35 yum; spores 5-6
X 9 mm.
Chemistry: Protocetraric and usnic acids, skyrin.
Type collection: Exfoliating granite outcrops in karoo, 78.6 km §S of
Sutherland on Hwy R354, elev. ca 900 m, Cape Province, S. Africa, Grid
8220 DC, M. E. Hale 73017, 29 Jan. 1986 (US, holotype; PRE, isotype).
This adnate conspersoid lichen may be related to X. dichromatica
(Hale) Hale, which is more loosely adnate and contains purplish
anthraquinone pigments. Xanthoparmelia tumidosa Hale (below) has convex,
more closely adnate lobes and lacks skyrin.
Xanthoparmelia subruginosa Hale, sp. nov. Fig. 52
Thallus laxe adnatus, saxicola, 6-10 cm latus, lobis sublinearibus,
elongatis, 0.5-1.5 mm latis, superne emaculatus vel sparse maculatus,
isidiis sorediisque destitutus, subtus planus, rugosus, niger, rhizinis
nullis.
Thallus loosely adnate on flat rocks, 6-10 cm broad, bright
yellowish green; lobes sublinear, dichotomously branched, 0.5-1.5 mm
wide, divaricate to subimbricate; upper surface continuous, shiny,
emaculate or weakly maculate on some older lobes, transversely cracked
with age; isidia and soredia absent; lower surface plane, smooth to
rugose, black, the tips with a yellowish rim, rhizines absent.
Pycnidia and apothecia absent.
Chemistry: Stictic, constictic and usnic acids.
Type collection: Flat sandstone exposures back from cliff, Oribi
Gorge Nature Reserve, elev. 300 m, Natal, S. Africa, Grid 3030 CB, M. E.
Hale 74042, 3 Feb. 1986 (US, holotype; PRE, isotype).
Another nonisidiate stictic acid-containing species which lacks
rhizines, X. albomaculata Hale, is generally more adnate, has shorter,
more subirregular lobes, a coriaceous thallus and distinct maculae on the
surface. It occurs at higher elevations in eastern Transvaal. Xantho-
parmelia suberadicata (des Abb.) Hale from Madagaascar has narrower, more
elongate, divaricate lobes, usually not more than 1 mm wide.
Additional specimens examined. S. Africa. Natal: Same locality as
the holotype, Hale 74018, 74019 (US).
603
Xanthoparmelia surrogata Hale, sp. nov. Fig. 53
Thallus adnatus, saxicola, 6-10 cm latus, coriaceus, lobis sub-
linearibus, 1-2 mm latis, superne emaculatus, isidiis sorediisque
destitutus, subtus planus, modice rhizinosus.
Thallus adnate to loosely adnate on rock, rather leathery, 6-10 cm
broad, dull yellowish green; lobes sublinear-elongate, 1-2 mm wide,
separate to subimbricate; surface continuous, shiny, emaculate, becoming
finely fissured toward the center; isidia and soredia absent; lower part
of medulla orange red; lower surface plane, black, shiny, moderately
rhizinate, the rhizines black, 0.5-1 mm long.
Pycnidia common; conidia bifusiform, 0.5 X 5-6 pm. Apothecia not
found.
Chemistry: Salazinic and usnic acids, skyrin.
Type collection: Large flat Table Mountain sandstone boulder in
Fynbos, trail above Woodhead Reservoir (to Platteklip Gorge), Table
Mountain Nature Reserve, elev. 800 m, Cape Province, S. Africa, Grid 3318
CD, M. E. Hale 72084, 23 Jan. 1986 (US, holotype; PRE, isotype).
Xanthoparmelia surrogata differs from X. luminosa (Elix) Hale, the
only other species with this chemistry and a black lower surface, in
having narrower, sublinear lobes and a leathery texture. Actually X.
luminosa occurs at two localities in southwestern Cape Province.
Xanthoparmelia tablensis Hale, Nash and Elix, sp. nov. Fig. 54
Thallus laxe adnatus, saxicola, 6-10 cm latus, lobis sublinearibus,
0.6-1.5 mm latis, superne emaculatus, isidiis sorediisque destitutus,
subtus planus, niger, modice rhizinosus.
Thallus loosely adnate on rocks, 6-10 cm broad, bright yellowish
green; lobes sublinear, elongated and little branched, 0.6-1.5 mm wide,
becoming laciniate, imbricate; upper surface continuous, emaculate;
isidia and soredia lacking; lower surface plane, black, sparsely to
moderately rhizinate, the rhizines black, 0.5-1 mm long, simple.
Pycnidia poorly developed; conidia bifusiform, 0.5 X 5-6 pm.
Apothecia adnate to substipitate, 1-3 mm in diameter; hymenium 45-50 pm;
spores 6-7 X 9-11 wm (data from Nash s.n.).
Chemistry: Hypoprotocetraric, 4-O-demethylnotatic and usnic acids
and several unidentified minor components.
Type collection: Large sandstone ledge in dense Fynbos, trail from
Platteklip Gorge to Woodhead Reservoir, Table Mountain Nature Reserve,
elev. 1000 m, Cape Province, S. Africa, Grid 3318 CD, M. E. Hale 72033,
23 Jan. 1986 (US. Holotype).
The only other loosely adnate, hypoprotocetraric acid-containing
species is X. hypoprotocetrarica (Kurok. & Elix) Hale, a common
effigurate—maculate lichen in S. Africa and Australia. The present
species is externally close to X. tasmanica (Hook. & Tayl.) Hale, which
is a larger lichen with salazinic acid. It has only been found on Table
Mountain.
Additional specimen examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 72036 (US); Devil's Peak, Table Mountain,
Grid 3318 CD, Nash s.n. (ASU).
Xanthoparmelia terricola Hale, Nash and Elix, sp. nov. Fig. 55
Thallus laxe adnatus, terricola, 6-15 cm latus, lobis sublinearibus,
1-1.5 mm latis, superne emaculatus, isidiis sorediisque destitutus,
subtus planus, brunneus, modice rhizinosus.
604
Thallus adnate to loosely attached on rocks, pebbles and soil, often
with the same thallus extending from soil to adjacent rocks, 6-15 cm
broad, rather firm, yellowish green but darkening with age at the center;
lobes sublinear, 1-1.5 mm wide, dichotomously branched, contiguous to
imbricate; upper surface continuous, shiny, emaculate, transversely
cracked and somewhat rugose with age; isidia and soredia absent; lower
surface plane, brown to dark brown, moderately rhizinate, the rhizines
brown, 0.5-1 mm long, simple or furcate with white tips when young.
Pycnidia common; conidia bifusiform, 0.5 X 5-6 pm. Apothecia
commonly developed, adnate, 3-8 mm in diameter; hymenium 40-45 pm;
spores 5-6 X 9-10 pm.
Chemistry: Salazinic, consalazinic, norstictic (trace),
protocetraric (trace) and usnic acids (determined by T. Nash, J. Elix and
J. Johnston).
Type collection: On soil and pebbles in area of flat sandstone
exposures, top of Brandwagsrot, Golden Gate National Park, elev. ca 1900
m, Orange Free State, S. Africa, Grid 2828 BC, M. E. Hale 74036, 11 Feb.
1986 (US, holotype; PRE, isotype).
This is the commonest soil-inhabiting lichen at higher elevations in
the Drakensberg escarpment. It has rather uniform, sublinear lobes with
a brown to dark brown lower surface, characters which distinguish it from
X. taractica (Kremplh.) Hale.
Additional specimens examined. S. Africa. Orange Free State: Same
locality as the holotype, Hale, 74000, 74005, 74007, 74008, 74009, 74010,
74026, 74037, 74050 (US). Transvaal: Long Tom Pass on Hwy R37, Grid 2430
BA, Hale 72046 (US); Mt. Sheba Nature Reserve, Grid 2430 DC, Hale 72050
(US), Nash 23417, 23418 (ASU). Lesotho: Sani Pass, Grid 2929 CB, Hale
74004, 74006, 74013, 74028, 74033. Natal: Witsieshoek Pass, Grid 2828 DD,
Hale 74011, 74012, 74014, 74016, 74025, 74027 (US).
Xanthoparmelia thamnolica Hale, sp. nov. Fig. 56
Thallus adnatus, saxicola, 5-8 cm latus, lobis subirregularibus,
brevibus, 1.3-2.5 mm latis, superne emaculatus, isidiatus, isidiis
cylindricis, subtus planus, niger, modice rhizinosus.
Thallus adnate on rocks, easily breaking apart, 5-8 cm broad, dark
yellowish green; lobes subirregular to sublinear, 1.3-2.5 mm wide,
imbricate; upper surface continuous, shiny, emaculate, isidiate, the
isidia becoming dense with age, cylindrical, simple to branched, 0.08-0.1
mm in diameter, 0.1-0.3 mm high; lower surface plane, black with a dark
brown zone at the tips, moderately to densely rhizinate, the rhizines
black, 0.5-2 mm long, simple to furcate.
Pycnidia and apothecia not seen.
Chemistry: Thamnolic and usnic acids.
Type collection: On large sandstone boulders in pasture, 25 km §S of
Clanwilliam along Hwy N7, elev. ca 300 m, Cape Province, S. Africa, Grid
3218 DD, M. E. Hale 72034, 25 Jan. 1986 (US, holotype; PRE, isotype).
Only one other Xanthoparmelia species contains thamnolic acid
(always with squamatic acid, lacking in X. thamnolica), X. cedrus-montana
Brusse, a member of the effigurate-maculate X. hypoleia group.
Superficially X. thamnolica would seem to be a rather large X. conspersa
(Ach.) Hale, a common species with stictic acid, which also occurs in S.
Africa. This species is highly restricted to the Clanwilliam area in
southern Namaqualand.
Additional specimens examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 72040 (US); 36.4 km §S of Clanwilliam,
Grid 3218 DD, Hale 72025, 72039 (US); Witelskloof, Grid 3218 BD, Brusse
605
Figures 53-56. Holotypes of Xanthoparmelia: 53, X. surrogata; 54, X.
tablensis; 55, X. terricola; 56, X. thamnolica. Scale = mm.
606
3092 (PRE); Versveld Pass, Grid 3218 DC, Brusse 2937 (PRE); 7 km W of
Tulbagh, Grid 3319 AC, Brusse 2714 (PRE); 32.7 km ESE of Citrusdal, Grid
3219 CA, Hale 72042 (US).
Xanthoparmelia toninioides Hale, sp. nov. Fig. 57
Thallus adnatus, saxicola, 4-6 cm latus, lobis subirregularibus, 1-2
mm latis, centrum versus valde bullatis, superne emaculatus, pruinosus,
isidiis sorediisque destitutus, subtus planus, ambitu niger, centro
pallide brunneus, modice rhizinosus.
Thallus adnate to tightly adnate on rocks, 4-6 cm broad, light
yellow green; lobes subirregular, 1-2 mm wide, short, contiguous,
becoming strongly bullate at the center; upper surface continuous, dull,
becoming densely pruinose; isidia and soredia absent; lower surface plane
to concave, blackish toward the tips but brown to pale brown toward the
center, moderately to densely rhizinate, the rhizines black or brown,
0.3-0.5 mm long.
Pycnidia and apothecia not found.
Chemistry: Hypoprotocetraric, 4-O-demethylnotatic and usnic acids.
Type collection: Sloping granite domes in Karoo, 1 km east of
Springbok on north side of Hwy R64, elev. 1100 m, Cape Province, S.
Africa, Grid 2917 DB, M. E. Hale 72086, 27 Jan. 1986 (US, holotype; PRE,
isotype).
The conspicuous bullate folds of thallus are filled with a
flocculent medulla. It is related to X. encrustans (Hale) Hale, a
subcrustose species, and probably confined to upper Namaqualand.
Additional specimen examined. S. Africa. Cape Province: Same
locality as the holotype, Hale 72086 (US).
Xanthoparmelia transvaalensis Hale, Nash and Elix, sp. nov. Fig. 58
Thallus adnatus, saxicola, 6-8 cm latus, lobis subirregularibus, 1-2
mm latis, margine laciniatis, superne emaculatus, isidiatus, isidiis
globosis, densis, subtus planus, nigricans, modice rhizinosus.
Thallus adnate on rock, 6-8 cm broad, darkish yellow green; lobes
subirregular to sublinear, 1-2 mm wide, imbricate with small marginal
laciniae ca 0.5 mm wide; upper surface continuous, shiny, emaculate,
isidiate, isidia dense and crowded, globose, basally constricted, mostly
unbranched, breaking off apically, 0.15-0.18 mm in diameter, 0.15-0.2 mm
high; lower surface plane, brown to black, sparsely to moderately
rhizinate, the rhizines brown or blackening, 0.5-1 mm long, simple.
Pycnidia lacking. Apothecia rare, substipitate, 2-5 mm in diameter;
hymenium 45 wm; spores poorly developed, 5 X 7-8 ym (data from Nash
23441).
Chemistry: Dehydroconstipatic (major), pertusaric (major),
constipatic (trace), protoconstipatic (trace) and usnic acids with other
unidentified fatty acids (same as in X. subdecipiens (Vain.) Hale)
(determined by J. Johnston).
Type collection: On sandstone outcrop in pasture, summit of Long Tom
Pass, Farm De Kuiten 205, S side of Hwy R37, elev. 2140 m, Lydenburg,
Transvaal, S. Africa, Grid 2430 BA, M. E. Hale 72088, 16 Jan. 1986 (US,
holotype; PRE, isotype).
This species has the same chemistry and black lower surface as X.
subnigra Hale, another species collected so far only at Long Tom Pass in
the Drakensbergs and perhaps representing the nonisidiate morphotype.
Additional specimens examined. S. Africa. Same locality as the
holotype, Hale 72027 (US), Nash 23441 (ASU).
607
Seeeerseans
HEEL
Figures 57-60. Holotypes of Xanthoparmelia: 57, X. toninioides; 58, X.
transvaalensis; 59, X. treurensis; 60, X. tumidosa. Scale = mm.
608
Xanthoparmelia treurensis Hale, Nash and Elix, sp. nov. Fig. 59
Thallus adnatus vel laxe adnatus, coriaceus, saxicola, 6-8 cm latus,
lobis sublinearibus, 2-5 mm latis, superne sparse maculatus, isidiatus,
isidiis cylindricis, subtus planus, rugosus, niger, rhizinis nullis.
Thallus adnate to loosely adnate on rocks, firm, 6-8 cm broad,
darkish yellowish green; lobes sublinear, 2-5 mm wide, contiguous; upper
surface continuous to weakly or strongly maculate in older parts, shiny,
transversely cracked, isidiate, the isidia moderately developed,
cylindrical, simple to coralloid branched, 0.15-0.2 mm in diameter, 0.5-2
mm high; lower surface plane, rugose, rhizines absent or very sparsely
developed.
Pycnidia and apothecia lacking.
Chemistry: Stictic, constictic, cryptostictic, norstictic (trace),
connorstictic (trace), constipatic and usnic acids (determined by T.
Nash, J. Elix and J. Johnston).
Type collection: On sandstone in coppice area, Treur River near
Bourkes Luck, elev. 1200 m, Pilgrims Rest, Transvaal, S. Africa, Grid
2430 DB, M. E. Hale 72037, 14 Jan. 1986 (US, holotype; PRE, isotype).
This species appears to be the isidiate morph of X. albomaculata
Hale, which has more distinct maculae. It is known only from two
localities in Transvaal but has probably been overlooked in the
escarpment area.
Additional specimens examined. S. Africa. Transvaal: Same locality
as the holotype, Hale 72044, 72048, 72049 (US), Sipman 19912 (B); Lejuma
Summit, Soutpansberg Mountains, Grid 2329 AB, Sipman 19873 (B).
Xanthoparmelia tumidosa Hale, sp. nov. Fig. 60
Thallus adnatus, saxicola, 3-6 cm latus, lobis sublinearibus,
brevibus, 1.5-2.0 mm latis, centrum versus congestis, bullatis, superne
emaculatus, convexus, isidiis sorediisque destitutus, subtus planus,
brunneus, modice rhizinosus.
Thallus adnate on rocks, leathery, 3-6 cm broad, bright yellowish
green; lobes short, sublinear, 1.5-2.0 mm wide, little branched,
contiguous but becoming crowded and bullate at the center; upper surface
continuous, dull, emaculate, convex—bullate; isidia and soredia absent;
lower surface plane to somewhat grooved, pale to darker brown, sparsely
to moderately rhizinate, the rhizines pale brown, ca 0.5 mm long, coarse,
unbranched.
Pycnidia common; conidia bifusiform, 0.5 X 5-6 pm. Apothecia
numerous, adnate, 1.5-2.5 mm in diameter, the rim rolling inward;
hymenium 55-60 um; spores 5-6 X 9-10 mm.
Chemistry: Protocetraric and usnic acids.
Type collection: Large exposed sandstone ledge in pasture, 1 km §S of
Nuwerus on Hwy R363, E of N7, elev. ca 500 m, Cape Province, S. Africa,
Grid 3118 AB, M. E. Hale 72027, 26 Jan. 1986 (US, holotype; PRE,
isotype).
The coriaceous, convex, adnate, almost inflated lobes distinguish
this Namaqualand-southwestern Cape lichen. There are no close relatives
with protocetraric acid.
Additional specimens examined. S. Africa. Cape Province: 1 km E of
Springbok, Grid 2917 DB, Hale 72047 (US); 8.8 km E of Garies, Grid 3018
AC, Hale 73108, 73110 (US); 44 km N of Vanrhynsdorp, Grid 3118 BC, Hale
73010, 73109, 73112 (US); 19.5 km NNW of Vanrhynsdorp, Grid 3118 BC, Hale
73111 (US); Kruisrivier, Grid 3321 BB, Hale 73011 (US); Seven Weeks
Poort, Grid 3321 AD, Brusse 3587 (PRE).
609
Xanthoparmelia viridis Hale, sp. nov. Fig. 61
Thallus laxe adnatus, saxicola, 6-10 cm latus, lobis subirregular—
ibus, 1.2-2.5 mm latis, superne maculatus, isidiis sorediisque destitu-
tus, subtus planus, brunneus vel nigricans, modice rhizinosus.
Thallus adnate to loosely adnate on rock, 6-10 cm broad, greenish
yellow; lobes subirregular to sublinear, 1.2-2.5 mm wide, imbricate,
irregularly laciniate; upper surface faintly to distinctly maculate,
shiny; isidia and soredia absent; lower surface plane, brown but black-
ening at the center, uniformly moderately rhizinate, the rhizines black,
1-1.5 mm long, simple.
Pycnidia common; conidia bifusiform, 0.5 X 6-7 um. Apothecia
numerous, adnate, 2-4 mm in diameter; hymenium 40-45 mum; spores 5-7 X
9-10 mm.
Chemistry: Fumarprotocetraric, succinprotocetraric and usnic acids.
Type collection: Schist ledges on hillside, Karoo Botanical Garden,
Worcester, elev. 300 m, Cape Province, S. Africa, Grid 3319 CC, M. E.
Hale 74023, 20 Jan. 1986 (US, holotype; PRE, isotype).
There is only one closely related species with fumarprotocetraric
acid, X. nuwarensis Hale (above), a Namaqualand lichen which is brown
below. Xanthoparmelia viridis may represent a chemotype of X. synestia
(Stirt.) Hale, a more widely distributed species in southwestern Cape
Province with salazinic acid.
Additional specimen examined. S. Africa. Cape Province: Kirstenbosch
Botanical Garden, Grid 3318 DC, Hale 72103 (US).
Xanthoparmelia wesselsii Hale, sp. nov. Fig. 62
Thallus laxe adnatus, terricola, tenuis, 4-7 cm latus, lobis
linearibus, elongatis, 0.5-1.5 mm latis, vix contrictis, apice
subteretibus, superne emaculatus, isidiis sorediisque destitutus, subtus
planus, brunneus, sparse rhizinosus.
Thallus loosely adnate on soil and humus, easily breaking apart, 4-7
cm broad; lobes linear, elongate, 0.5-1.5 mm wide, sometimes weakly
constricted, dichotomously branched, separate to divaricately imbricate,
the ultimate branches becoming terete, subascending; upper surface
continuous, shiny, emaculate; isidia and soredia absent; lower surface
plane to somewhat convoluted, shiny, light to darker brown in flaring
parts of lobes, interrupted by concolorous constricted parts, sparsely
rhizinate, the rhizines brown to dark brown, 0.5-1 mm long.
Pycnidia not well developed; conidia not seen. Apothecia adnate,
3-4 mm in diameter; hymenium 42-45 jim; spores 5-6 X 9-10 ym.
Chemistry: Salazinic, consalazinic and usnic acids.
Type collection: On soil among small sandstone outcrops in pasture,
summit of Long Tom Pass on S side of Hwy R37, Farm De Kuiten, Lydenburg,
elev. 2140 m, Transvaal, S. Africa, Grid 2430 BA, M. E. Hale 72043, 16
Jan. 1986 (US, holotype; PRE, isotype).
This soil-inhabiting lichen from higher elevations in the
Drakensbergs has a lower surface consisting of flattened, flaring light
brown parts a few mm long and intervening constricted areas concolorous
with the upper surface. Another terricolous lichen in the Drakensbergs,
X. terricola Hale, Nash & Elix, has broader, more appressed lobes. This
species is named in honor of Dirk Wessels.
Additional specimens examined. S. Africa. Transvaal: Same locality
as the holotype, Hale 72017, 72018, 72019 (US), Sipman 200781 (B); Mt.
Anderson, Leighton 3272 (BOL); 14 km N of Dulistroom, Grid 2530 AC,
Drews CH4359 (PRE).
610
PiLLLIit
Figures 61, 62. Holotypes of Xanthoparmelia: 61, X. viridis; 62, X.
wesselsii. Scale = mm.
MYCOTAXON
WOMEN. pps, 6141-616 October-December 1986
a a ear a ae ee eee tie i
NEW SPECIES OF RELICINA (LICHENIZED ASCOMYCOTINA)
FROM AUSTRALASIA
JOHN A. ELIX and JEN JOHNSTON
Department of Chemistry, The Faculties, Australian National University, GPO Box 4,
Canberra, ACT, 2601, Australia
ABSTRACT : The species, Relicina agglutinata Elix&Johnston and Relicina
niuginiensis Elix&Johnston from Papua New Guinea, Relicina conglutinata
Elix&Johnston, Relicina demethylbarbatica Elix&Johnston and Relicina
filsonii Elix&Johnston from Australia, and Relicia fijiensis Elix&Johnston
from Fiji are described as new. In addition the new combination, Relicina
butleri (Elix) Elix&Johnston is made.
Introdution
The lichen genus Relicina Hale, a segregate of Parmelia Acharius sens. lat., was
erected by Hale (Hale 1974) to accommodate those species with narrow, adnate lobes,
marginal bulbate cilia and usnic acid in the upper cortex. Relicina is characterised further
by an upper cortex consisting of a basic palisade plectenchyma with a pored epicortex
(Hale 1973) and short, bifusiform conidia. In a subsequent monograph Hale (Hale
1975) considered Relicina to comprise of 24 species with the major centre of distribution
being in south-east Asia and Indonesia. At that time a total of seven species were known
to occur in the Australasian region and a further twelve have been recognised since ( Elix
& Stevens 1979; Kashiwadani 1975; Kurokawa 1979; Stevens 1981), but we now wish
to describe a further six new species.
Throughout the present work chemical constituents were identified by thin layer
chromatography ( Culberson 1972; Culberson & Johnson 1982), high performance
liquid chromatography ( Lumbsch & Elix 1985) and comparison with authentic samples.
Relicina agglutinata Elix & Johnston, sp. nov. Fig.1
Species thallo ut in Relicina sublanea simili sed ab hac specie acido barbatico et acido
4-O-demethylbarbatico continens differt.
Type. Papua New Guinea. Milne Bay Province. On tree branch in lowland forest,
Sagarai, between Wegulani and Badili Rivers, 10°24'S. 150°17'E, 40m, R.Kumei 136
pr.p., 16.1.1985 ; CBG-holotype.
Thallus foliose, corticolous, loosely adnate on bark, pale yellow-green to
yellow-brown, to 8cm in diameter; /obes continuous and sublinear to linear-elongate and
separate, 1.0-2.0(-3.0)mm wide, barely or not imbricate, dichotomously or
subdichotomously divided, the margins bulbate ciliate, cilia sparse, black, inflated and
becoming globose at the base but paler and often developing agglutinated hyphae
towards the apices, up to 2.0mm long. Upper surface more or less convex, continuous,
dull, weakly maculate, lacking soredia and isidia; medulla white; lower surface pale tan,
primary rhizines moderate, scattered, black or blackening, but with a thick mass of fine,
pale tan, densely branched or agglutinated secondary rhizines developing from the
primary rhizines or directly from the lower surface. Apothecia common, adnate, to 2mm
in diameter, disc more or less flat or shallowly concave, pale brown to cinnamon-brown
; the exciple ecoronate, margin dentate: asci 8-spored, spores simple, colourless,
612
ellipsoid, 5-7 x 4-5 tum.
Chemistry. Thallus K-, medulla K-, C-, KC+ yellow, P- ; containing usnic acid,
barbatic acid (major), 4-O-demethylbarbatic acid (minor), obtusatic acid (trace),
norobtusatic acid (trace), protocetraric acid (trace).
Morphologically this new species closely resembles Relicina sublanea (Kurok.) Hale.
Both species are characterised by the pale lower surface with fine, densely branched,
pale, agglutinated secondary rhizines arising from the lower surface and primary
rhizines, the separate, linear-elongate lobes and the ecoronate apothecia. However these
two species are distinguished readily by their medullary chemistry, with R. sublanea
reacting P+ orange-red due to the presence of protocetraric acid, whereas R. agglutinata
gives a P- reaction (barbatic acid and 4-O-demethylbarbatic acid). Although these two
species are sympatric in Papua New Guinea, R. sublanea has a more extensive and more
northerly distribution, being known from Indonesia, Malaysia and the Philippines (Hale
1975). The only other Relicina with comparable chemistry, R. incongrua Hale, is
distinguished readily by the jet black, sparsely rhizinate lower surface and the
coproduction of diffractaic acid.
Specimens Examined:
Papua New Guinea. Morobe Province. On large Castanopsis in Castanopsis forest on
south-west facing slope, Nauti Logging Area, Upper Watut River, 10km south-west of
Bulolo, 7°16'S, 146°36'E, 1400m, H.Streimann 17063, 3.iii.1982 (CBG8201190,
LAE).
Relicina conglutinata Elix & Johnston, sp. nov. Fig.2
Species thallo ut in Relicina circumnodata simili sed ab hac specie acido
succinprotocetrarico et acido fumarprotocetrarico continens differt.
Type. Australia. Queensland. On mangroves in tidal flats, 17°01'S, 145°46'E, 1m,
J.A.Elix 17616 & H.Streimann, 8.vii.1984 ; CBG-holotype.
Thallus foliose, corticolous, closely adnate on bark, yellow-green, to Scm in
diameter; /obes sublinear-elongate, 0.8-1.5mm wide, separate, sparingly imbricate,
dichotomously divided, the margins bulbate ciliate, cilia moderate to sparse, commonly
vestigial and subglobose to globose, black, then black and inflated at the base but paler
towards the apices, up to 0.3mm long. Upper surface plane to convex, continuous,
shiny at the apices but dull on the older lobes, weakly maculate, moderately to densely
isidiate, isidia subcylindrical, short, irregularly inflated, simple or rarely sparingly
branched; medulla white; lower surface pale tan, primary rhizines moderate to dense,
mottled black and pale tan, with slender secondary rhizines developing from the primary
rhizines or directly from the lower surface, secondary rhizines densely branched and
becoming conglutinated, secondary rhizines pale, concolorous with the lower surface.
Apothecia not seen.
Chemistry. Thallus K-, medulla K-, C-, KC-, P+ brick red; containing usnic acid,
succinprotocetraric acid (major), fumarprotocetraric acid (minor), protocetraric acid
(trace).
This new species closely resembles the common species, Relicina circumnodata
(Nyl.) Hale as they have comparable thalli and lobe configurations, produce cylindrical
isidia on the upper surface and have pale conglutinated secondary rhizines below. In
general the cilia of R. conglutinata are more sparse and poorly developed than those of
R. circumnodata with many vestigial bulbae not developing into true cilia, but these two
species are distinguished most readily chemically. Thus the major medullary substance in
R. circumnodata is protocetraric acid, while R. conglutinata produces
succinprotocetraric acid and fumarprotocetraric acid. R. conglutinata appears to be the
isidiate counterpart of Relicina ramosissima (Kurok.) Hale. At present R. conglutinata
is known only from the type locality.
Relicina demethylbarbatica Elix & Johnston, sp. nov. Fig.3
Species thallo ut in Relicina abstrusa simili sed ab hac specie acido
4-O-demethylbarbatico (maximo) et acido barbatico (minimo) continens differt.
Type. Australia. Queensland. On felled trees along the rainforest margin, Flaggy Creek,
613
Figures 1-6. New species of Relicina: 1, R.agglutinata (holotype in CBG); 2,
R.conglutinata (holotype in CBG); 3, R.demethylbarbatica (holotype in CBG); 4,
R fijiensis (holotype in CBG); 5, R.filsonii (holotype in MEL); R.niuginiensis (holotype
in CBG). Scale in mm.
614
Black Mountain Road, 9km west of Kuranda, 16°47'S. 145°36'E, 420m, J.A.Elix
17663 & H.Streimann, 7.vii.1984 ; CBG-holotype.
Thallus foliose, corticolous, closely adnate on bark, yellow-green, to 8cm in
diameter; lobes short, sublinear-elongate, 0.3-1.5mm wide, crowded but sparingly
imbricate, the margins bulbate-ciliate; cilia numerous, moderately inflated, up to 0.3mm
long. Upper surface plane, continuous, shiny, emaculate, moderately to densely isidiate;
isidia cylindrical, short, simple or rarely sparingly branched; medulla white; lower
surface jet black, densely rhizinate, rhizines black, simple. Apothecia not seen.
Chemistry. Thallus K-, medulla K-, C-, KC+ yellow, P-; containing usnic acid,
4-O-demethylbarbatic acid (major), barbatic acid (minor).
This new species closely resembles the common species, Relicina abstrusa (Vainio)
Hale as they have comparable thalli and lobe configurations and both have simple,
cylindrical isidia on the upper surface. Generally thalli of R. demethylbarbatica are more
diminutive, but these two species are distinguished most reliably chemically. Thus the
major medullary substance in R. abstrusa is norstictic acid which gives K+ yellow
turning red, C-, P+ orange medullary reactions, while the medulla of R.
demethylbarbatica reacts K-, C-, P-, KC+ yellow due to the presence of
4-O-demethylbarbatic acid. The unique ‘chemistry distinguishes this new siecle from ali
known Relicina .
Specimens Examined:
Australia. Queensland. In "dry" rainforest on Acacia , slopes of Black Mountain, 25km
north-west of Kuranda, 16°40'S, 145°29'E, 500m, J.A.Elix 17492 & H.Streimann,
7.vii.1984 (ANUC); on bark, logging area in rainforest, end of Clohesy River Road,
a.. 16km south-east of Kennedy Hwy., west of Cairns, 800m, M.E.Hale, s.n.,
8.viii.1983 (US); on bark, logging area in rainforest, near end of Black Mountain Road,
33km west-north-west of Kuranda, 800m, M.E.Hale 64085, 20.viii.1983 (US).
Relicina fijiensis Elix & Johnston, sp. nov. Fig.4
Species thallo ut in Relicina samoensis simili sed ab hac specie acido incognito
continens differt.
Type. Fiji. Viti Levu. On trees along the roadside in regrowth forest, Nasori Highlands,
Nadi-Sigatoka Road, 3.6km west of Vanturu Dam turnoff, J.A.Elix 15116,
26.viii.1983 ; CBG-holotype.
Thallus foliose, corticolous, closely adnate on bark, yellow-green, to Scm in
diameter; lobes short, sublinear, 0.3-1.0mm wide, crowded but sparingly imbricate, the
margins bulbate-ciliate; cilia conspicuous, bulbae globose, cilia up to 1.0mm long, rarely
branched. Upper surface plane, slightly shiny, emaculate, lacking soredia and isidia;
medulla white; lower surface jet black, sparsely rhizinate, rhizines black, simple or
sparingly branched. Apothecia common, adnate, to 1.5mm in diameter, disc more or
less flat, pale brown to brown; the exciple coronate, amphithecium retrosely rhizinate;
asci 8-spored, spores simple, colourless, broad-ellipsoid, 5.5-7 x 3.5-4.5mm.
Chemistry. Thallus K-, medulla K+ pale yellow, C-, KC-, P-; containing usnic acid
and hirtifructic acid [Rp in solvents A/B/C (Culberson 1972): 50, 31, 38; solvent B'
(Culberson & Johnson 1982): 35; develops as a pale orange spot after spraying with
sulfuric acid and heating].
This new species would appear to be closely related to Relicina samoensis (Zahlbr.)
Hale as they have comparable thalli, coronate apothecia, a black lower surface and lack
isidia. However the thalli of R. fijiensis are compact, with more or less contiguous
lobes, dense cilia and retrose apothecial rhizines, whereas R. samoensis has separate
lobes with moderately dense cilia and retrose rhizines. Even so these two species are
distinguished most reliably chemically. Thus the major medullary substance in R.
samoensis is echinocarpic acid while R. fijiensis produces hirtifructic acid. The latter
substance was first recognized in Relicina hirtifructa (Kurok.) Streim. although the
structure of this compound remains unknown. This new species differs morphologically
from R. hirtifructa since the latter species has much more robust thalli, broader lobes
(1.5-3.0mm) and ecoronate apothecia.
Specimen Examined:
615
Fiji. Viti Levu. On tree in regrowth forest, Nasori Highlands, Nadi-Sigatoka Road,
13km east of the Vanturu Dam turnoff, J.A.Elix 15224, 27.viii.1983 (ANUC).
Relicina filsonii Elix & Johnston, sp. nov. Fig.5
Species thallo ut in Relicina limbata simili sed ab hac specie acido norstictico
(maximo), acido connorstictico (minimo) et acido hyposalazinico (vestigio) continens
differt.
Type. Australia. New South Wales. On tree, Kerripit River, Gloucester Tops, R.Filson
11174, 15.1.1969 ; MEL 1028512-holotype.
Thallus foliose, corticolous, adnate on bark, yellow-green to yellow-brown on
storage, to 5cm in diameter; lobes linear-elongate, contiguous centrally to separate at the
periphery, 1.5-3.0mm wide, not imbricate, subdichotomously divided, the margins
bulbate-ciliate; cilia dense and conspicuous, black, to 2.0mm long. Upper surfate plane
to weakly convex, continuous, shiny at the apices, weakly maculate, lacking soredia and
isidia; medulla white; lower surface brown to dark brown, rhizines dense, brown to
black, simple or occasionally dichotomously branched. Apothecia common, adnate, to
- 3mm in diameter, disc more or less flat or shallowly concave, cinnamon-brown; the
exciple ecoronate, amphithecium retrorsely rhizinate: asci 8-spored, spores simple,
colourless, ellipsoid, 7-9 x 4-6 um.
Chemistry. Thallus K-, medulla K+ yellow-red, C-, KC-, P+ orange; containing
usnic acid, norstictic acid (major), connorstictic acid (minor), hyposalazinic acid (trace).
Morphologically this new species closely resembles Relicina limbata (Laurer) Hale.
Both species are characterised by the brown lower surface with broad, linear-elongate
lobes which lack isidia, and the ecoronate apothecia. However these two species are
distinguished readily by their medullary chemistry: R. limbata reacts K+ yellow due to
the predominance of stictic acid (with minor quantities of norstictic, constictic and
cryptostictic acids), whereas R. filsonii gives a K+ yellow then red reaction due to the
predominance of norstictic acid (with minor quantities of connorstictic and hyposalazinic
acids). Although these two species are sympatric in New South Wales, R.limbata has a
more extensive distribution, being known from Tasmania to north Queensland. At
present this new species is known only from the type locality. The only other Relicina
with comparable chemistry and morphology, R.subabstrusa Hale, is distinguished by
the jet black lower surface.
Relicina niuginiensis Elix & Johnston, sp. nov. Fig.6
Species thallo ut in Relicina relicinula simili sed ab hac specie thallo elobulato et acido
gyrophorico et acido incognito continens differt.
Type. Papua New Guinea. Morobe Province. On upper branches of large tree on
Castanopsis and Dipterocarpaceae-dominated ridge, Herzog Mountains, 15km
west-south-west of Lae, 6°45'S, 146°51'E, 760m, H.Streimann 10932, 13.1.1981 ;
CBG-holotype
Thallus foliose, corticolous, closely adnate on bark, yellow-green, to 3cm in
diameter; lobes sublinear-elongate, 0.3-1.0mm wide, separate, not imbricate,
dichotomously divided, the margins bulbate-ciliate; cilia very dense and conspicuous, up
to 1.0mm long, simple or often branched. Upper surface plane, shiny, emaculate,
lacking soredia and isidia; medulla white; lower surface jet black, sparsely rhizinate,
rhizines black, simple or sparingly branched. Apothecia common, adnate, to 1.5mm in
diameter, disc more or less flat, pale brown to brown; the exciple coronate,
amphithecium retrorsely rhizinate; asci 8-spored, spores simple, colourless,
broad-ellipsoid, 5-5.5 x 3.5-4.5mm.
Chemistry. Thallus K-, medulla K-, C-, KC-, P+ yellow; containing usnic acid,
gyrophoric acid (trace), hirtifructic acid (major), unknown (trace).
This new species would appear to be closely related to Relicina relicinula (Mill.Arg.)
Hale since both species have narrow, sublinear-elongate, dichotomously divided lobes
which are well separated at the periphery. These two species can, however, be
distinguished both morphologically and chemically. R. relicinula typically is lobulate
616
with continuous, crowded central lobes and contains only usnic acid and pale unknowns
while R. niuginiensis is elobulate with well separated lobes throughout and contains
usnic acid, gyrophoric acid and hirtifructic acid. This species also could be confused
with R. fijiensis since they are similar morphologically and chemically. However the
lobes of R. fijiensis are shorter, more compact and crowded and this species contains
only hirtifructic acid and usnic acid. R. niuginiensis is distinguished by the more
elongated, well separated lobes and the presence of additional gyrophoric acid and an
unknown substance (which appears to be related to the compounds present in Relicina
butleri (Elix) Elix & Johnston).
Specimen Examined:
Papua New Guinea. Morobe Province. On Pometia trunk, in Endospermum medullosum
and Pometia pinnata dominated forest on slope, Buso River, 26km east-north-east of
Lae, 6°43'S, 147°14'E, 160m, H.Streimann 11621 & T.Umba, 16.i.1981 (CBG
8105180).
New Combination
Relicina butleri (Elix) Elix & Johnston comb.nov.
Basionym: Parmelia butleri Elix, Aust.J.Bot., 27, 876 (1979).
ACKNOWLEDGEMENTS
We thank the Australian National University for generous financial support of this
project through the Faculties Research Fund. We thank Dr.M.E.Hale for his
cooperation, advice and assistance in obtaining critical type material. We also wish to
thank Mr.I.A.Fox and the staff of the Science Photographic Unit, ANU, for preparing
the photographs, Mr.D.Verdon for checking the Latin descriptions and Dr. G.A.Jenkins
who determined the chemistry of many specimens. We thank the following colleagues
for making collections available: Mr.R.Filson (MEL), Dr.M.E.Hale (US),
Mr.P.Lambley (UPNG), Mr.H.Streimann (CBG), Dr.S.Kurokawa (TNS).
LITERATURE CITED
Culberson, C.F. (1972). Improved conditions and new data for the identification of
lichen products by a standardized thin-layer chromatographic method.
J. Chromatogr., 72, 113-125.
Culberson, C.F. and Johnson, A. (1982). Substitution of methyl tert.-butyl ether for
diethyl ether in the standardized thin-layer chromatographic method for lichen
products. J. Chromatogr., 238, 483-487.
Elix, J.A. and Stevens, G.N. (1979). New species of Parmelia (lichens) from
Australia. Aust. J. Bot., 27, 873-883.
Hale, M.E.Jr. (1973). Fine structure of the cortex in the lichen family Parmeliaceae
viewed with a scanning electron microscope. Smithsonian Contrib. Bot., 10, 1-92.
Hale, M.E.Jr. (1974). Bulbothrix, Parmelina, Relicina, and Xanthoparmelia, four new
genera in the Parmeliaceae (Lichenes). Phytologia, 28, 334-339.
Hale, M.E.Jr. (1975). A monograph of the lichen genus Relicina (Parmeliaceae).
Smithsonian Contrib. Bot., 26, 1-32.
Kashiwadani, H. (1975). Enumeration of Anaptychiae and Parmeliae of Papua New
Guinea. In Otani, Y. : Reports on the cryptogams in Papua New Guinea
(Natn.Sc.Mus., Tokyo), 75-85.
Kurokawa, S. (1979). Enumeration of species of Parmelia in Papua New Guinea. In
Kurokawa, S. : Studies on cryptogams of Papua New Guinea (Academia Scientific
Book Inc., Tokyo), 125-150.
Lumbsch, H.T. and Elix, J.A. (1985). A new species of the lichen genus Diploschistes
from Australia. Pl. Syst. Evol., 150, 275-279.
Stevens, G.N. (1981). The macrolichen flora on mangroves of Hinchinbrook Island,
Queensland. Proc. R. Soc. Qd., 92, 75-84.
AUTHOR INDEX, VOLUME TWENTY-SEVEN
ABDULLAH, SAMIR K., see UDAGAWA & AL.
AGERER, R., Studies on ectomycorrhizae III. Mycorrhizae formed by four fungi in the
genera Lactarius and Russula on spruce 1-59
ANON., Notice: Guide to Sanctioned Fungal Names 404
AWAO, TAKEYOSHI, see UDAGAWA & AL.
BAGYANARAYANA, G., & U. BRAUN, A new species of the genus Oidium from India 61-
62
BEN-ZE'EV, ISRAEL S., Notes on Entomophthorales (Zygomycotina) collected by T.
Petch: II. Erynia ellisiana sp. nov., non Erynia forficulae (Giard), comb. nov.,
pathogens of Forficulidae (Dermaptera) 263-269
BONDE, M. R., see MICALES & AL.
BORSE, B. D., see HYDE & BORSE
BRAUN, U., see BAGYANARAYANA & BRAUN
BRODO, IRWIN M., A new species of the lichen genus Sulcaria (Ascomycotina,
Alectoriaceae) from California 113-117
BRUSSE, FRANKLIN A., Five new species of Parmelia (Lichenes, Parmeliaceae) from
southern Africa 183-192
BRUSSE, FRANKLIN A., Four new effigurate-crustose species of Parmelia (Lichenes,
Parmeliaceae) from southern Africa 237-245
BURDSALL, HAROLD H., JR., Platygloea acanthophysa, a new species with single
sterigmate basidia and acanthophyses 499-502
CANDOUSSAU, FRANCOISE, Récoltes de Biscogniauxia dennisii (Pyrenomycetes,
Ascomycetes) dans le sud-ouest de la France 555-562
COURTECUISSE, REGIS, Notes de nomenclature concernant les Hyménomycétes; IV. Sur
quelques épithétes spécifiques préoccupés. 3. 127-145
ELIX, JOHN A., & JEN JOHNSTON, New species of Relicinia (lichenized Ascomycotina)
from Australasia 611-616
ELIX, JOHN A., JEN JOHNSTON, & DOUGLAS VERDON, Canoparmelia, Paraparmelia and
Relicinopsis. Three new genera in the Parmeliaceae (lichenized Ascomycotina) 271-
282
ERIKSSON, OVE, Lahmia Korber (= Parkerella A. Funk) a misinterpreted genus with
isolated position 347-360
ERIKSSON, OVE, see YUE & ERIKSSON
ERIKSSON, OVE, & JING-ZHU YUE, Bertiella (Sacc.) Sacc. & Sydow a synonym of
Massarina Sacc. 247-253
FRANK, HARALD M., Macrochemical color reactions of macromycetes II. Chemical
properties and systematic position of Bondarzewia mesenterica 503-506
FUNK, A., Two new discomycetes on Pinus 283-288
GJRUM, HALVOR B., East African rusts (Uredinales), mainly from Uganda 5. On
families belonging to Gamopetalae 507-550
HALE, MASON E., JR., New species of the lichen genus Xanthoparmelia from southern
Africa (Ascomycotina: Parmeliaceae) 563-610
HALLENBERG, NILS, Cultural studies in Tubulicrinis and Xenasmatella (Corticiaceae,
Basidiomycetes) 361-375
HANLIN,R. T., & R. A. SAUNDERS, Confirmation of Ascosphaera apis in Georgia 551-
554
HO, H. H., & S. C. JONG, A comparison between Phytophthora cryptogea and P.
drechsleri 289-319
HUMBER, RICHARD A., see ROMBACH & AL.
HYDE, K. D., & B. D. BORSE, Marine fungi from Seychelles. VI. Massarina velataspora,
a new marine Ascomycotina from mangrove wood 161-167
JOHNSTON, JEN, see ELIX & JOHNSTON
JOHNSTON, JEN, see ELIX & AL. ti :
JONES, E. B. GARETH, Digitatispora lignicola, anew marine lignicolous Basidiomycotina
155-159
618
JONG, S. C., see HO & JONG
KOSKE, R. E., & CHRISTOPHER WALKER, Species of Scutellospora (Endogonaceae) with
smooth-walled spores from maritime sand dunes: two new species and a redescription
of the spores ofScutellospora pellucida and Scutellospora calospora 219-235
KUNWAR, I. K., J. B. MANANDHAR, & J. B. SINCLAIR, Badarisama sojae, anew bulbil-
forming genus and species 119-125
a M . PAGE, Basidiomycetes that decay Gambel oak in southwestern Colorado: III.
5-345.
MANANDHAR, J. B., see KUNWAR & AL.
MICALES, J. A., M. R. BONDE, & G. L. PETERSON, The use of isozyme analysis in
fungal taxonomy and genetics 405-449
MILLER, STEVEN L., Hypogeous fungi from the southeastern United States I. The genus
Rhizopogon 193-218
NAGARKAR, M. B., P. K. SETHY, & P. G. PATWARDHAN, Materials for a lichen flora of
the Andaman Inslands —I 71-82
ONOFRI, SILVANO, see ZUCCONI & ONOFRI
PATWARDHAN, P. G., see NAGARKAR & AL.
PETERSON, G. L., see MICALES & AL.
PERINI, CLAUDIA, A new Chlamydomyces species from Italy 255-261
REYNOLDS, DON R., Foliicolous ascomycetes 7. Phylogenetic systematics of the
Capnodiaceae 377-403
ROBERTS, DONALD W., see ROMBACH & AL.
ROMBACH, MICHIEL C., R ICHARD A. HUMBER, & DONALD W. ROBERTS, Metarhizium
flavoviride var. minus, var. nov., a pathogen of plant- and leafhoppers on rice in the
Philippines and Solomon Islands 87-92
RYVARDEN, L., Tremellostereum (Tremellaceae) nov. gen. 321-323
SANDERS, F. E., see WALKER & SANDERS
SAUNDERS, R. A., see HANLIN & SAUNDERS
SCHIPPER, M. A. A., Hyphomucor, a new genus in the Mucorales for Mucor assamensis
83-86
SETHY, P. K., see NAGARKAR & AL.
SIMMONS, EMORY G., Gibbago, a new phaeodictyoconidial genus of hyphomycetes 107-
111
SINCLAIR, J. B., see KUNWAR & AL.
TSUBOUCHI, HARUO, see UDAGAWA & TSUBOUCHI
UDAGAWA, SHUN-ICHI, & HARUO TSUBOUCHI, Coniochaetidium mirabile, a new
ascomycete isolated from salted food 63-69
UDAGAWA, SHUN-ICHI, TAKEYOSHI AWAO, & SAMIR K. ABDULLAH,
Thermophymatospora, a new thermotolerant genus of basidiomycetous hyphomycetes
99-106
VERDON, DOUGLAS, see ELIX & AL.
WALKER, CHRISTOPHER, see KOSKE & WALKER
WALKER, CHRISTOPHER, & F. E. SANDERS, Taxonomic concepts in the Endogonaceae:
II. The separation of Scutellospora gen. nov. from Gigaspora Gerd. & Trappe 169-
182
WEBER, WILLIAM A., The lichen flora of the Galapagos Islands, Ecuador 451-497
YUE, JING-ZHU, see ERIKSSON & YUE
YUE, JING-ZHU, & OVE ERIKSSON, Studies on Chinese ascomycetes. 3. Astrosphaeriella
lageniformis 93-98
ZUCCONI, LAURA, & SILVANO ONOFRI, Two new dematiaceous hyphomycetes from
tropical forest litter 147-153
619
INDEX TO FUNGOUS AND LICHEN TAXA,
VOLUME TWENTY-SEVEN
This index includes the names of genera, infrageneric taxa, species, and
infraspecific taxa. New names are italicized, as are the page numbers on
which such new taxa are proposed. New names of suprageneric taxa appear
in italic CAPITALS.
Absidia 84
Acarospora chlorophana 458; schleicheri 459
Acaulospora 229, 231
Acremoniella atra 259
Acrogenotheca 378, 381, 384, 386, 388, 398, 399;
elegans 382
Actinomucor 84
Aecidium 511; acanthacearum 535; aspiliae 507,
508, 544; barleriae 527; celosiae 544;
cephalandrae 520; clerodendri 514; conyzae-
pinnatilobatae 512; cookeanum 546; crasso-
cephali 510, 511; cynanchi 508, 545; dichro-
cephalae 509, 545; dissotides 536; dubiosum
509, 547; echuyaénse 507, 509, 510, 545;
emiliae 510, 511; englerianum 544; flavidum
510, 546; fluggeae 547; formosanum 510, 511;
gynurae 510, 511, 545; gynurae-cernuae 510,
$11; haplocarphae 507, 511, 512, 545; hart-
wegiae 544; helichrysi 509; hoffmannii 512;
justicia 533; macowanianum 512, 545; mariani-
raciborskii 511; metapense 509; mitracarpi 520;
ocimi 531, 546; ornithogaleum 546; plectranthi
$12, 513, 547; plectranthicola 513; popowiae
546; pycnostachydis 513, 547; ranunculacearum
547; torae 544; verononiae-podocomae 513, 548
Agaricus 407, 505; bisporus 409; brunnescens 436;
campestris 436; cyaneus 138, 139; incanus v
polychrous 132; undulatellus 142; virescens 128-
131
Aigialus grandis 161; parvus 161
Alectoria 113, 114, 276; imshaugii 114; sarmen-
tosa 481, 488
Aleurodiscus cerrusatus 344
Allantoparmelia 274
Alpova 202; subg Antridium 202; klikae 202
Alternaria 107, 110, 111, 120; alternata 119, 120,
122; helianthi 110; leucanthemi 110
Amphinema byssoides 342
Anaptychia circinalis 470; comosa 470; coral-
lophora 470; leucomelaena 488; podocarpa 470,
488; stellata 470; f verrucifera 470; tropica v
antillarum 470
Angiopsora wiehei 517, 523
Aniptodera mangrovii 161
Antennulariella 378, 381, 384, 386, 388, 390,
393, 394, 398, 399
‘ANTENNULARIELLOIDEAE’ [nom. prov.] 398
Anthracothecium leucostomum 459; ochraceofla-
vum 459; pyrenuloides 459
Armillaria mellea 325, 326, 341
Arthonia 488; cinnabarina 460, 488; gregaria 460,
488; nivea 488; platyspilea 459, 460; tumidula
460
Arthrothelium galapagoense 459-461; spiloma-
toides 460
Arthrorhaphis grisea 356
Ascidium massalongoi 80; xanthostromiza 78
Ascosphaera apis 551-553
Aspergillus 255-260
Aspidophyllum fugiens 460, 469, 486
Astrosphaeriella 93, 97; aosimensis 97; lagenifor-
mis 93, 94, 97, 98
Athelia bombycina 343; coprophila 343
Bacidia 459; apiahica 460; insularis 459, 460;
luteola 460; millegrana 460; rubella 460;
subleporina 461
Badarisama 119, 120, 123, 124; sojae 119-121,
122, 124
Bertia subg Bertiella 249; macrospora 248-251;
moriformis v latispora 247; parasitica 249
Bertiella 247, 249, 250; macrospora 247, 253;
polyspora 250
Biatriospora marina 161
Biscogniauxia 555, 557; dennisii 555-561; margi-
nata 557; repanda 557
Bondarzewia 503; mesenterica 503, 505; montana
503
Borrera leucomelos 454
Botryobasidium botryosum 343
Bremia lactucae 437
Brigantiaea leucoxantha 461
Brooksia 398
Bryoria 113, 114, 276; sect Implexae 114; furcel-
lata 114; kuemmerleana 114; smithii 114; spira-
lifera 114
Buellia 459, 488, 491; canescens 466; dejungens
461; galapagona 459, 461; glaziouana 451, 493;
lauricassiae 461; mamillana 451, 461, 493;
modesta 461; parasema 461; straminea 459, 461,
462, 488, 490; umbrina 488; xanthinula 488
Bulbothrix 276; laevigatula 462
Burgoa 119, 124; nigra 124; verzuoliana 120, 124
Byssoloma subdiscordans 462, 474, 485;
tricholomum 462
Byssoporia terrestris 44, 45, 47, 48; v aurantiaca
47; v sartoryi 47
Calopisma byrsonimae 462
Caloplaca 463; byrsonimae 462; camptidia 462;
cirrochroa 462; elegans 463, 488; floridana 463;
isidiosa 463, 488, 489; muelleri 463, 488, 491;
620
[Caloplaca] murorum 489; rugulosa 463, 489
Calosphaeria 355
Canoparmelia 271-277; adspersa 278; amazonica
278; aptata 278; caribaea 274, 278; carneo-
pruinata 274, 278; caroliniana 274, 278;
cinerascens 278; concrescens 278; corrugativa
278; crozalsiana 274, 278; cryptochlorophaea
278; ecaperata 276, 278; epileuca 278; eruptens
278; inhaminensis 278; inornata 278; martin-
icana 278; nairobiensis 278; norsticticata 278;
raunkiaeri 278; salicinifera 274, 279; schelpei
279; scrobicularis 279; somaliensis 279; sub-
tiliacea 279; texana 274, 277, 279; zambiensis
276, 279; zimbabwensis 279
Capnodium 378, 381, 382, 386, 388, 394, 397,
399; walteri 380, 384
Caryosporella rhizophorae 161
Ceraceomyces serpens 325, 327, 343
Ceratobasidium cornigerum 326, 327, 343
Ceratocladium 149
Ceriporia excelsa 341; reticulata 341; tarda 341
Ceriporiopsis mucida 341
Chiodecton effusum 463; farinaceum 463; myrti-
cola 463; sanguineum 469, 489
Chlamydomyces 255, 259, 260; diffusus 259;
palmarum 259, 260; sympodialis 255, 257, 258,
260
Chrysosporium 103
Chrysothrix candelaris 463, 466
Circinotrichum 147, 149; britannicum 149; fertile
149, 150; flagelliforme 147-149; papakurae 149;
rigidum 149, 150
Cladia 453; aggregata 463
Cladina 453, 464; confusa 463, 489; sandstedei 464
Cladobotryum 63, 67
Cladonia 463, 494; balfourii 464, 489; cariosa 465;
ceratophylla 464; chlorophaea 464, 489; didyma
464; fimbriata 489; furcata f adspersa 489;
galapagosensis 463, 464, 489; macilenta ssp
theiophila 464; pityrea 465, 489; polia 463, 464,
489; pycnoclada 489; sphacelata 465; subcariosa
465; subsquamosa 465; subulata 465; sympho-
riza 465
Claudopus 140; subg Claudopus 140; subg
Paraleptonia 140; cyaneus 140; cyanomelaenus
140
Claussenomyces 287, 288; dacrymycetoideus 287;
olivaceus 287; pini 283, 284, 287; prasinulus
287; pseudotsugae 287
Clavaria 323
Clitopilus flavidus 140, 141
Coccocarpia erythroxyli 466; palmicola 465, 466,
490; pellita v parmelioides 466
Coccoidea 355
Coccotrema granulatum 490
Coenogonium 489
Coleosporium 538; clematidis 544; mitteri 538
Collema furfuraceum 466
Colletotrichum 437
Conidiobolus 264
Coniochaetidium 63, 67; mirabile 63, 65-68;
ostreum 63, 66, 67
Coprinus cinereus 103, 409; delicatulus 103
Cora pavonia 466, 489
Cortinarius subg Leprocybe 49; hochstetteri 132
Corynella 287
Cronartium coleosporioides 284
Crumenulopsis 288; lacrimiformia 283, 284, 286,
287; pinicola 288
Cryphonectria 436
Dacrymyces minor 340
Dendrostilbella 283
Dictyonema guadelupense 466, 489; montanum
466, 489; sericeum 466, 489
Didymella 250
Dietelia impatientis 546
Digitatispora lignicola 155, 157, 158; marina 155,
157, 158
Dimerella lutea 466
Diploicea canescens 463, 466
Diploschistes actinostomus 466, 489; bryophilus
467; muscorum 467; scruposus 467, 490
Dirina approximata 467, 489; herrei 467, 489
Dirinaria 489-491; aegialita 467, 491; applanata
467, 468, 491; caesiopicta 467, 491; confluens
467; leopoldii 468, 491
Disporotrichum dimorphosporum 105
Eccilia 144; subg Nolanea 141; subg Omphalopsis
144; sect Endochromonema 141; subsect Cheilo-
cystidiata 141; subsect Icterina 141; flavida 141,
142; mexicana 142; tenuipes 143, 144
Echinoplaca 462
Embellisia 107, 111
Empusa forficulae 263; v major 263, 264, 266-268
Endocarpon pusillum 468
Endogone 171; calospora 180; heterogama 180
Endophyllum 514; macowanii 514; striatosporum
514, 547; superficiale 514, 544
Endothia gyrosa 436; parasitica 436
Entoloma 127, 128, 132, 140, 143, 144; subg
Leptonia 133, 139; subg Paludocybe 133; sect
Albidicaules 133; sect Leptonia 139; sect Paludo-
cybe 133; sect Staurospori 133; aeruginosum
128-132; albidum 132, 133; allochroum 139;
canum 132; cinerascens 132; convexum 133,
134; cyaneum 138-140; dichroum 139; egonii
127, 133; flavidum 140, 141; flavoides 127, 141,
142; gracile 133; guzmanii 127, 143; hebes 144;
hesleri 133, 134; hochstetteri 132; largentii 127,
134, 135, 138; v alnicola 127, 136, 137; v
badiodorsum 127, 135, 136; v farinaceum 127,
137, 138; leptopus 144; mexicanum 142, 143;
omphaliaeformis 144; porphyrophaeum 138;
subaeruginosum 127, 128; subcaeruleum 142;
tenuipes 144; tjallingiorum 139; velatum 139;
violaceum 138; virescens 127, 129, 131
Entomophthora 264, 407; forficulae 263, 264, 266-
268
Erioderma pusillum 468
Erynia 263, 266, 268; subg Furia 268; subg
Zoophthora 269; ellisiana 263, 264, 266-268;
forficulae 263, 268, 269; suturalis 266
Euantennaria 378, 381, 382, 386, 388, 390, 391,
393, 394, 396, 398, 399
Everniastrum vexans 468, 490
Exidia glandulosa 339, 340
Fibulomyces mutabilis 343
Flavoparmelia 272-276
Flavopunctelia 272, 274
Fomitopsis pinicola 370
Fracchiaea heterogena 250
Fusarium solani 409
Gibbago 107, 108, 111; trianthemae 107, 108,
110, 111
Gigaspora 169-172, 174, 177-179, 226; albida 179;
alborosea 180; aurigloba 180; calospora 171,
174, 175, 177, 179, 180; candida 179; coralloidea
181; decipiens 179; dipapillosa 181; erythropa
181; gigantea 171, 177, 179; gilmorei 181;
gregaria 181; heterogama 170, 171, 175, 177,
178, 180; margarita 170-172, 178, 179; minuta
180; nigra 174, 175, 181; pellucida 181; persica
181; reticulata 169, 181; rosea 179, 226;
savannicola 180; tricalypta 180; verrucosa 181
Gloeophyllum sepiarium 326, 328, 342; trabeum
328, 342
Gloeoporus dichrous 321
Glyphus 469; afzelii 469; caesiella 469; cicatricosa
468; desquamescens 469; striatula 469; subamy-
lacea 469
Graphina virginea 468
Graphium 67
Gyalectidium filicinum 469
Gymnoderma 494
Gymnopilus sapineus 341
Gyrothrix 149; pediculata 149
Haematomma puniceum 469
Halocyphina villosa 158
Halosarpheia abonnis 161; ratnagiriensis 161
Hamaspora engleriana 547; longissima 547
Hapalopilus nidulans 341
Hemileia canthii 514, 515, 544; holstii 515, 547;
scholzii 515, 544; vastatrix 515, 545; woodii
516, 547
Herpothallon sanguineum 469, 489
Heterodermia albicans 469; antillarum 469, 470;
comosa 470, 488; corallophora 470; lepidota
470; leucomela 454, 470, 471, 488, 491;
lutescens 471
Hygrophorus cyaneus 132; hochstetteri 132
Hymenochaete arida 344
Hyphoderma anasaziense 325, 328, 329, 344;
medioburiense 329, 330, 345; praetermissum 345
Hyphodontia 364, 365, 374; arguta 342; crustosa
339, 342; subalutacea 342
Hyphomucor 83, 84; assamensis 83, 84-86
Hypogymnia mollis 113
Hypotrachyna costaricensis 471; microblasta 471
Hypoxylon 557
Idriella 150; fertilis 150
621
Incrustoporia nivea 341
Inonotus andersonii 339, 342; dryophilus 341
Itersonilia perplexans 499, 501
Kavinia himantia 331, 342
Kirschsteinia polyspora 250
Lactarius 1, 14, 46, 47, 503; deliciosus 10, 12, 14;
v picinus 10; deterrimus 1, 2, 4, 6, 8, 10, 12,
14, 21, 46, SO, 51; griseus 50; maculatus 50;
obscuratus 49; picinus 1, 2, 8, 10, 15, 16, 18,
20) 21,0123; 46°50; rufus. 45, 46, 517.53:
salmonicolor 14; sanguifluus 10, 14, 51
Laeticorticium simplicibasidium 499, 501
Lahmia 347, 348, 352-357; fueistingii 356; kunzei
347-353, 355-358; picea 356
LAHMIACEAE 357
LAHMIALES 357
Lahmiomyces 356
Lecanora 490; atra 485; caesiorubella 114; ssp
glaucomodes 471; floridana 463; glaucovirens
489; leprosa 471, 472; leptoplaca 242; pseudo-
pinguis 459, 461, 471; punicea 469; sulfurescens
471
Lecidea 472; aeruginosa 472; chilena 472; coarctata
485; glaziouana 493; granifera 471, 472, 490;
granulosa 485-486; oreinoides 472, 488
Lentinellus cochleatus 340
Lentinus edodes 409, 436
Lepolichen coccophorus 490; colobinus 490
Lepraria candelaris 463
Leprocaulon microscopicum 472; tenellum 472
Leptogium cyanescens 472; isidiosellum 472;
marginellum 473; milligranum 473; punctulatum
473
Leptonia aeruginosa 128; albida 132, 133;
cinerascens 132; convexa 133-135; v alnicola
136; v badiodorsa 134, 135; v farinacea 138;
cyanea 138; mexicana 142; subcaerulea 142;
undulatella 142
Leptoniella acericola 139; aeruginosa 128; albida
132-134; mexicana 142
Leptosphaeria avicenniae 166; coccodes 251;
contecta 166; senegalensis 250
Leptotrema 72, 489; bahianum 74; compunctum
474; elachistoteron 72, 82; mastoideum 490;
nuwarense 71, 74, 82; pertusarioides 71, 74, 82;
reclusum 72, 74, 82; wightii 474
Limacinia 378, 381, 382, 386, 388, 390, 394,
396-399; fernandeziana 382, 384
‘LIMACINIOIDEAE’ {nom. prov.] 397
Limacinula 378, 381, 382, 386, 388, 393, 394,
397, 399; anomala 384
Lobaria dissecta 473
Lopadium foliicola 473, 485; fuscum 473;
puiggarii 473 :
Lophiosphaera 250; beckhausii 252
Lophiostoma 247, 250, 251, 253; angustilabrum
251; arundinis 249, 251; semiliberum 249, 251;
subcorticale 249-251
Lophiotrema 250
Lophodermium juniperinum 353
622
Massarina 161, 165, 166, 247, 250, 251, 253;
aquatica 165, 166; coccodes 247, 251, 253;
cystophorae 165, 166; eburnea 250; macrospora
247, 248, 250-253; ‘sp. 1' 166; ‘sp. 2' 166;
velataspora 161-166
Melampsora euphorbiae 545, 547
Melanaspicilia 493
Melanelia 274
Melanotheca cruenta 473
Metacapnodium 378
Metarhizium 87, 91; album 88, 91; anisopliae 87,
88, 90; v anisopliae 88; v majus 88; flavoviride
87-90, 92; v flavoviride 87, 89-91; v minus 87,
89-92
Metasphaeria 250, 251; sepincola 251
Minimedusa 119, 124
Mortierella 84
Mucor 83, 84; assamensis 83, 84; 'tax. sp. II' 86
Myceliophthora 99, 103
Mycogone 259; cervina 259; rufa 259
Myriotrema compunctum 474; nuwarense 74;
olivaceum 76, 474; wightii 474
Neofuscelia 274
Neovossia horrida 437
Neurospora 407
Nia vibrissa 158
Niebla 474; ceruchoides 474
Nimbospora bipolaris 166
Nitschkia broomeiana 250
Nodulisporium 557
Nolanea tenuipes 144
Normandina pulchella 474
Nummularia 557; dennisii 556; discreta 557
Ocellularia 72; allosporoides 75, 82; berkeleyana
75, 78, 82; cavata 79; diacida 75, 82; glaucula
76; glaziovii 78; micropora 76; olivacea 71, 76,
82; pycnophragmia 76, 82; terebrans 76, 82;
wandoorensis 71, 78, 82; xanthostromiza 71, 78,
82
Ochrolechia pallescens 474
Oidium 61, 62; abelmoschii 62; candicans 343;
pavoniae 61, 62; schmiedeknechtii 62; urenae 62
Ophiocapnocoma 378, 394
Pannaria molybdaea 466, 490; pannosa 474
Panus rudis 340
Papulaspora 119; atra 124; pannosa 120, 124;
parasiticum 119
Paraparmelia 271-277, 279; adusta 279; alabamen-
sis 279; annexa 279; arcana 279; astricta 279;
basutoensis 279; capnoides 276, 280; cerussata
280; condyloides 280; conranensis 280; fumigata
276, 280; interposita 276, 280; ischnoides 280;
lecanoracea 280; lithophila 280; lithophiloides
280; molybdiza 280; mongaensis 280; murina
280; nana 280; neoquintaria 280; numinbahensis
280; owariensis 280; prolata 280; pustulescens
280; rodriguesiana 280; rupicola 281; schistacea
276, 281; scotophylla 279, 281; spodochroa 281;
subtortula 281; tortula 281; vanderbylii 281;
violacea 281; xanthomelaena 281
Parathalle 356; fueistingii 356
Parkerella 347, 348, 354, 356; populi 347, 348,
S5023577 308
Parmelia 183, 185, 237, 240, 271, 272, 453, 611;
subg Cyclocheila 271; subg Parmelia 271; sect
Cyclocheila 271; sect Xanthoparmelia 184, 187,
188; ‘group’ Parmeliae 237; ‘group’ Xanthopar-
meliae 242; adspersa 278; adusta 279; ala-
bamensis 279; amazonica 278; americana 468;
annexa 279; aptata 278; arcana 279; astricta 185,
240, 279; basutoensis 279; butleri 616; camt-
schadalis 468, 490; capnoides 280; caribaea 278;
Carneopruinata 278; caroliniana 278; cerussata
280; cinerascens 278; concrescens 278; condy-
loides 280; conformata 475; conranensis 280;
corrugativa 278; costaricensis 471; crinita 475;
cristifera 475, 476; crozalsiana 278; cryptochloro-
phaea 278; dilatata 475; dominicana 475; eborina
475; ecaperata 278; endosulphurea 475; epileuca
278; eruptens 278; eximia 183, 184, 190; fausta
183, 185, 186, 190, 240; fumigata 280; hypo-
tropa 476; ianthina 576; inhaminensis 278;
inornata 278; insignis 183, 185, 186, 187, 190,
240; insipida 237, 238, 239, 243; interposita
280; intertexta 281; inuncta 183, 187, 190;
irregularis 188; ischnoides 280; laevigatula 462;
latissima 475, 477, 490; lecanoracea 280, 577;
leptoplaca 237, 238, 242; leucosemotheta 476;
lithophila 280; lithophiloides 188, 280; malac-
censis 282; martinicana 278; microblasta 471;
molybdiza 185, 280; mongaensis 280; murina
280; nairobiensis 278; nana 280; neoquintaria
280; nimbicola 237, 238, 240, 243; norsticticata
278; numinbahensis 280; owariensis 280;
peralbida 476; perlata 475, 490; praesorediosa
476; prolata 280; pudens 183, 188, 190, 240;
pustulescens 280; rahengensis 282; ralla 237,
240-242, 244; raunkiaeri 278; reticulata 476;
rodriguesiana 280; rupicola 281; salacinifera 279;
schelpei 279; schenckiana 185; schistacea 281;
scotophylla 281; scrobicularis 279; somaliensis
279; soyauxii 490; sphaerospora 184; spodochroa
281; squamatica 237, 242, 244; subcrinita 476;
subdecipiens 187, 188; subramigera 487; sub-
tiliacea 279; subtortula 281; texana 279; tincto-
rum 476; tortula 281; unctula 187; vanderbylii
281; violacea 281; xanthina 476; xanthomelaena
190, 281; zambiensis 279; zimbabwensis 279
Parmeliella pannosa 474
Parmentaria astroidea 475
Parmotrema 272, 276; conformatum 475; crinitum
475; cristiferum 475, 490; dilatatum 475;
dominicanum 475; eborinum 475; endosulphu-
reum 475, 490; hypotropum 476; leucosemothe-
tum 476; peralbidum 476; praesorediosum 476;
reticulatum 476; subcrinitum 490; tinctorum
476, 490; ultralucens 476; xanthinum 477
Paxillus involutus 52
Peltigera didactyla 477; dolichorhiza 477; erumpens
[Peltigera erumpens]477; hazslinszkyi 477; spuria
477
Peltula 480; euploca 477
Peniophora nuda 334, 344; pseudoversicolor 344;
violaceolivida 344
Perenniporia medulla-panis 341; tenuis 342
Peronosclerospora 407, 437
Peronospora tabacina 437
Pertusaria 74, 477, 490; albinea 459, 477; bispora
477; colobina 490; pustulata 478
Phaeographis dendritica 478
Phaeomarasmius 339; erinaceus 338, 339, 341,
345
Phaeophyscia hispidula 478
Phaeotrema 72; lacteum 79, 82
Phakopsora 540; apoda 546; gossypii 545;
incompleta 546, 547; loudetiae 546; pachyrhizi
436, 548; setariae 547; vernoniae 516, 548
Phanerochaete arizonica 345; carnosa 345;
chrysosporium 103; sanguinea 331, 332, 345;
sordida 332, 333, 345; tuberculata 344; velutina
344
Phellinus ferruginosus 342
Phillipsiella atra 355
Phlebia albida 342; lilascens 343
Pholiota erinacea 339
Phragmocapnias 378, 381, 386, 388, 393, 394,
397, 399
Phyllopsora corallina 478
Physcia 479; adglutinata 490; aegialita 467, 491;
alba 491; albicans 469; aspera 467, 491; biziana
478; callosa 478; insularis 459, 478; lacinulata
479; leucomela 470, 491; mexicana 478, 479;
picta 467, 491; sorediosa 479
Physma byrsinum 479
Physopella africana 544; cameliae 546; hansfordii
544; hiratsukae 545; nyasalandica 517, 545; tecta
545; wiehei 517, 523, 545
Phytophthora 289, 314, 315, 407; cinnamomi 290,
292, 293, 296, 297, 299-304, 312-313; crypto-
gea 289-301, 303, 305-309, 312-315; drechsleri
289-295, 298, 299, 301-307, 309-315; erythro-
septica 314; himalayensis 314; infestans 408,
437; megasperma 315; nicotianae 290, 292, 293;
palmivora 290, 292, 295, 300, 302; richardiae
314; sinensis 303
Placodium isidiosum 463; murorum 463, 489
Placopsis cribellans 491
Platygloea 499, 500, 502; acanthophysa 499, 500-
502; laplata 325, 333, 334, 340; mycophila 499;
peniophorae 334; unicornis 501; unispora 499
Pleospora guadefroyi 166
Polyporus arcularius 339, 342
Poria terrestris 44
Porina 462
Porosphaerellopsis 355
Porosphaeria 355
Protoparmelia 237; badia 237
Protoparmeliopsis 237; muralis 237
Pseudocyphellaria 453; argyracea 479; aurata 454,
468, 479; crocata 480; mougeotiana v aurigera
480, 491; xantholoma 480
623
Pseudoparmelia 27 1-277; caperata 271-272; caroli-
niana 480; chapaadensis 271; cyphellata 271;
dahlii 281; hypomilta 271; intertexta 272; leuco-
xantha 480; scotophylla 272; sphaerospora 271;
texana 272
Psoroglaena cubensis 480
Ptychogaster 105
Puccinia 407, 521; absinthii 519; abutili 544, 548;
aecidiiformis 517, 546; aframomi 544; africana
518, 547; ambigua 522; andropogonicola 546;
andropogonis 545; anthriscicola 544; arenariae
$44, 547; artemisiae-afrae 507, 518, 519, 544;
artemisiae-sibiricae 519; arthraxonis-ciliaris 544;
aurata 518; bakoyana 520, 546; becii 520, 544;
borreriae 520; brachypodii v arrhenatheri 545; v
poae-nemoralis 547; calcitrapae 523; cardui-
pycnocephali 436; carduorum 436; carthami 436;
cenchri v africana 544, 546; cephalandrae 520,
521, 548; cephalandrae-indicae 521; chaerophylli
$44; chaseana 544; cnici-oleracei 519; collettiana
532; conclusa 545; coronata 545; crassocephali
507, 521, 545; cyathulicola 545; cynodontis 545;
cyperi-cristati 545; cyperi-tagetiformis 545;
desertorum 522, 545; dietelii 544; difformis 522,
545; digitariae-velutinae 545, 547; dimorpha 532;
dummeri 523, 545; duthiae 544; echinopsis 523;
emaculata 546; eritraeensis 544, 546; erlangeae
517, 523, 545; eucomi 546; exilis v hibisci 546;
fimbristylidis 545; flaccida 546; fragilispora 508;
fragosoana 546; fusispora 545, 547; garckiana
546; graminis 436; f sp tritici 408, 409;
guizotiae 524, 545; hansfordiana 524, 547;
hedbergii 524, 547; heterospora 547; holosericea
525, 546; hoslundiae 525, 546; hyparrheniae
546; hypoéstis 526; isoglossae 526, 546;
kalchbrenneri 509, 545; v kalchbrenneri 526; v
valida 526; keniensis 546; lantanae 526, 527,
544; lateritia 520, 528, 547; leonotidicola 528,
546; leptosperma 545; leucadis 529, 539, 546;
leveillei 545; levis v panici-sanguinalis 544; v
tricholaenae 547; lippiicola 529, 530, 546;
lippiivora 529; lithospermi 522; loudetiae 546;
macowanii 509; melanocephala 547; menthae
530, 547; morotorensis 545; nabugobo-ensis
547; nakanishikii 545, 547; natalensis 530, 546;
v evansii 529; necopina 530, 531, 547;
oahuensis 545; obscura 546; obscuripora 547;
ocimi 531; perlargonii-elgonensis 547; palargo-
nii-zonalis 546; pentadis-carneae 531, 547;
petitiana 544; peucedani-kerstenii 547; phyllo-
cladiae 544; pinarii 509; plectranthi 513;
polygoni-amphibii 546, 547; polysora 548;
pseudechinolaenae 547; pulvinata 523; recondita
409, 436, 544; rubiicola 531, 532, 547;
rubiivora 532; ruelliae 532, 533, 547; saniculae
547; schizocarphae 521; semiloculata 533, 546;
smilacis-kraussianae 547; sorghi 548; stobaeae v
stobaeae 534; v woodii 533, 534, 544; tanaceti
519; tetradeniae 507, 534, 535, 547; thunbergiae
533, 535, 544, 545, 547; thunbergiae-alatae 535;
tragiae 547; vernoniicola 536, 548; versicolor
546; violae 548; zorniae 548
624
Pucciniosira dissotidis 536, 545
Punctelia 272, 274
Pycnoporus cinnabarinus 339, 341
Pyrenula aurantiaca 480; cerina 480; nitida 475
Pyrgillus javanicus 480
Pyricularia 147, 152; oryzae 437; buloloensis 152;
fusispora 152; lauri 152; sphaerulata 147, 150-
152
Pyxine 491; berteriana 481; caesiopruinosa 481;
cocoes 481; connectens 481; endoleuca 491;
eschweileri 481; glebosa 462, 491; pringlei 481,
491
Ramalina 452; anceps 481, 482; ceruchoides 474;
complanata 482, 492; dasypoga 491; denticulata
482; farinacea 491; furcellata 482, 491; indica
492; interponens 492; linearis 482, 492;
peruviana 482; puiggari 492; sorediantha 482;
subasperata 492; subfraxinea 492; usnea 453,
481, 482, 488, 491, 492; usneoides 492
Ravenelia albiziae-zygiae 544; cassiaecola v
berkeleyi 544; hansfordii 544; indigoferae 546;
spegazziniana 105; tephrosiae 547
Relicina 274, 276, 611, 614, 615; abstrusa 612,
614; agglutinata 611, 613; butleri 611, 616;
circumnodata 612; conglutinata 611, 612, 613;
dimethylbarbatica 611, 612-614; fijiensis 611,
613, 614, 616; filsonii 611, 613, 615; hirti-
fructa 614; limbata 615; niuginiensis 611, 613,
615, 616; ramosissima 612; relicinula 615;
samoensis 614; subabstrusa 615; sublanea 611
Relicinopsis 271-277, 281; dahlii 281; intertexta
272, 281; malaccensis 282; rahengensis 282
Resupinatus applicatus 340
Rhizomucor 84
Rhizopogon 193, 196, 197, 199, 200, 203, 204,
206-208, 213-218; stirps Luteolus 202; abietis
193, 194, 204, 205, 208, 209, 217; atlanticus
193, 194, 214, 215, 217; brownii 205, 206;
cokeri 212, 214; couchii 193, 195, 196, 204,
205, 207, 209-212, 217, 218; evadens 193, 196,
210-212; v evadens 195, 211, 213, 217; fabri
193, 194, 199, 201, 216; fuscorubens 193, 195-
197, 216; hesleri 218; lowii 198, 200; luteolus
52, 201; maculatus 215; nigrescens 193, 195,
197, 198, 216; parasiticus 215; piceus 201;
roseolus 215; v intermedius 208; rubescens 193,
195, 202-205, 215, 216; separabilis 196; smithii
211; subaustralis 193, 195, 203-206, 208, 216;
succosus 193, 194, 198-201, 208, 216; v
mexicanus 201; superiorensis 198, 200; v
mexicanus 200, 201; truncatus 193, 194, 212-
215, 217; vinicolor 44, 47-49; vulgaris 193,
195, 204-212, 217, 218; v intermedius 208
Rhizopus 84
Rhodophyllus aeruginosis 129, 132; cyaneus 140;
holocyaneus 129-132; mexicanus 142, 143
Rhynchostoma 93, 96-98; lageniforme 93, 94, 97,
98; minutum 93, 95, 96; rubrocinctum 95, 96
Rhytidiella 347; baranyayi 347, 357; moriformis
347, 358
Rinodina 493; contiguella 493; mamillana 493;
thomae 493
Roccella 492; babingtonii 452, 482, 492; difficilis
492; galapagoensis 459, 483, 492; intricata 482;
peruensis 482; portentosa 482
Roccellina badia 483
Roccellodea nigerrima 492
Russula 1, 32, 34, 46-48, 503-505; sect Ingratae
34; sect Rigidae 40; adusta 50; atropurpurea 504;
caerulea 503; chloroides 504; cyanoxantha 504;
fellea 504, 505; fragilis 50; grisea 40, 42;
heterophylla 504; ochroleuca 1, 2, 25, 26, 29,
30, 32, 34, 40, 46, 48, 50, 504, 505; sardonia
50; vesca 40, 42; virescens 504; xerampelina 1,
2, 30, 32, 34-36, 38, 40, 42, 44-46, 48, 50 504
Sarrameana 353
Schistophoron tenue 483
Scorias 378, 381, 386, 388, 393, 394, 397, 399;
spongiosa 380, 382
Scutellospora 169, 171, 179, 219, 226, 228, 233,
234; alborosea 180, 226; aurigloba 180, 234;
calospora 180, 219, 222, 228, 231, 233, 234;
coralloidea 181; dipapillosa 181; erythropa 181;
fulgida 219, 221, 222; gilmorei 181, 222;
gregaria 181; heterogama 180; minuta 180; nigra
181; pellucida 181, 219, 222, 226, 228, 229,
231; persica 181; reticulata 181; savannicola 180,
224; tricalypta 180; verrucosa 181; weresubiae
219, 224-226, 228
Septobasidium 499, 501, 502
Seuratia 355
Sistotrema brinkmannii 343
Sphacelotheca reiliana 437
Sphaeria 250
Sphinctrina podocarpa 483
Spilonema revertens 483
Sporopodium leprieurii 484
Sporotrichum 105; azureum 105; pruinosum 103,
105; pulverulentum 103
Steccherinum fimbriatum 342
Stegobolus berkeleyanus 75
Stemphylium 107, 110
Stereocaulon 453, 494; apocalyptum 451, 494;
azulense 451, 459, 467, 484, 494; microcarpum
484; weberi 459, 484; wrightii 451, 494
Stereum 321; dichroum 321, 323; hirsutum 344
Sticta 453; aurata 454; filix 484; fuliginosa 484;
quercizans 492; sylvatica 484; weigelii 484, 492;
f peruviana 485
Strigopodia 378, 381, 382, 386, 388, 390, 393,
394, 396, 398, 399
Strigula 462; nitidula 485; subtilissima 485
Stropharia 138
Subulicystidium longisporum 335, 344
Suillus 45, 48, 50; luteus 47
Sulcaria 113-115; badia 114; isidiifera 113, 115,
117; sulcata 114; virens 114
Tapellaria 462; epiphylla 462, 485; nana 469, 485;
nigrata 485
Teloschistes chrysophthalmus 142; exilis 492;
flavicans 485, 493
Tephromela atra 485
Thaxteria 354
Thelotrema 72; allosporoides 75; depressum 80;
elachistoteron 72; glaucopallens 79, 82; guptei
71, 79, 82; lacteum 79; massalongoi 80, 82;
polillense 80, 82; pycnophragmium 76; reclusum
74; recurvum 71, 80, 82; terebrans 76
Thermomucor 84; indicae-seudaticae 86
Thermophymatospora 99, 100; fibuligera 99, 100,
102-104
Tilletia caries 437; controversa 437; foetida 437;
indica 408, 437
Tomentella ferruginea 343; molybdaea 335, 336,
343; pilosa 342; ramosissimum 343; subtestacea
336, 337, 342
Trapelia coarctata 485
Trapeliopsis granulosa 485
Trechispora farinacea 336, 337, 343; vaga 343
Trematosphaeria lignatilis 161, 166
Tremellostereum 321, 323; dichroum 321-323
Tricharia vulgaris 486
Trichobasis palmarum 259
Tricholomopsis platyphylla 337, 338, 341
Trichomerium 378, 381, 386, 388, 390, 391, 397,
399
Trichopeltheca 378, 381, 382, 386, 388, 390, 391,
396, 398, 399
Trichosphaeria 93
Triposporiopsis spinigera 378
Troposporella fumosa 358
Trypethelium eluteriae 486; mastoideum 486;
ochroleucum 486; tropicum 486
Tubeufia 356
Tubulicrinis 361, 369, 374; borealis 361, 363,
365, 369, 370; calothrix 344, 361, 363, 365,
369; glebulosus 364; gracillimus 361, 364, 365,
369; medius 361, 365, 368, 369; strangulatus
361, 363, 365, 370; subulatus 361, 365, 371
Ulocladium 107, 110
Uredo 512, 533, 540, 545; achyranthicola 544;
bidenticola 540; caricis-confertae 544;
clerodendricola 536; clerodendrina 536, 537, 544;
desmodii-ramosissimi 545; dissotidis 536;
ecteinanthi 538; fuirenae-strictae 545; humbertii
537, 545; hypoéstis-verticillaris 507, 537, 546;
isoglossae 507, 538, 546; kabaleensis 547;
kabanyoloensis 544, 550; leucadicola 507, 538,
546; mitteri 538; montis-elgonensis 544;
paederiae 507, 539, 546; palmarum 259;
palpigera 539, 540, 546; pycnostachydis 513;
ruelliae 532; scholzii 515; sporoboli-
pyramidalidis 547; stylosanthis 547; xyridis 548
Uromyces 542; appendiculatus 547; aspiliicola
540; bidenticola 540, 544; blainvilleae 540, 544;
clignyi 544, 546, 547; cluytiae 545; commelinae
544-546; crotalariae 545; dipcadi 545; eulophiae
545; euphorbiae 545; geranii 545; hobsonii 541,
546; hyparrheniae 546; hypoéstis 537;
625
indigoferae 546; ipomoeae 541, 546; keniensis
544; linearis 546; melantherae 542, 546;
notabilis 545; pegleriae 545; pseudarthriae 547;
rhynchosiae 547; sasaénsis 507, 542, 547;
setariae-italicae 544-547; striatus 546; tenuicutis
547; transversalis 545; valerianae 542; valerianae-
wallichii 542, 543; vignae 548; wiehei 547
Usnea 493; amabilis 493; antillarum 486;
arthroclada 487; barbata 493; ceratina 493;
Cladocarpa 487; longissima 493; paradoxa 487;
plicata 453, 493; rubescens 487; rubiginea 487,
493
Ustilago scitaminea 437
Xanthoparmelia 276, 277, 563, 565, 569, 571,
$74-576, 579-581, 583, 584, 587-589, 591-593,
595, 596, 599-601, 604, 605, 607, 610; acrita
563, 564, 565; adhaerens 582; africana 563, 564,
565; albomaculata 602, 608; aliphatica 563, 565,
566, 567, 600; australasica 568; austroafricana
598; barbellata 592; barklyensis 563, 565, 566;
calvinia 563, 567-570; capensis 563, 567, 569;
cedrus-montana 604; ceresensis 563, 568, 569;
chlorochroa 576; colorata 570; competita 563,
$67, 568-570; conspersa 568; constrictans 597;
crassilobata 563, 570, 571; cumberlandia 580;
denudata 563, 570, 571, 598; diacida 563, 571,
572; dichromatica 602; duplicata 563, 571, 572,
591; effigurata 563, 573, 575 586; encrustans
606; endochromatica 563, 574-576, 580, 598;
endomiltoides 576, 588; enteroxantha 563, 574,
575; epigaea 563, 573, 575, 576; equalis 563,
576, 579; esterhuyseniae 563, 577, 579; evernica
563, 567, 577, 579, 596; exornata 586;
globisidiosa 563, 578, 579; gyrophorica 563,
574, 578, 581; huachucensis 572; hybrida 563,
580, 581; hypoleia 573, 585, 604; hypo-
protocetrarica 603; hyporhytida 572; ianthina
563, 576; indumenica 563, 580, 581; kalbii 596;
karooensis 563, 581, 582, 592; lagunebergensis
563, 577, 582, 583, 596; lecanoracea 563, 577;
lesothoensis 563, 582, 583; leucostigma 574,
580; lividica 563, 583, 584, 592; lobulifera 563,
583, 584; luminosa 594, 603; minuta 563, 585,
587, 591; mollis 563, 585, 587; molliscula 577;
mougeotii 594; mougeotina 585; namakwa 563,
573, 586, 587; namaquensis 563, 586, 587;
neocongensis 592; neosynestica 563, 588, 589;
neotasmanica 563, 588, 589; nigropsoromifera
572; nuwarensis 563, 589, 590, 609; ochro-
pulchra 563, 589, 590; olifantensis 563, 591,
593; olivetorica 563, 574, 584, 591, 593;
oribensis 563, 592, 593; phaeophana 590;
probarbellata 563, 592-594; pustulifera 563, 594,
595; rubro-medulla 563, 594, 595; saniensis 563,
595, 596; schenckiana 572; serusiauxii 563, 577,
578, 582, 595, 596; simulans 563, 597, 599;
skyrinifera 563, 597, 599; stenosporonica 563,
574, 598, 599; subcolorata 563, 572, 597, 598,
599; subcrustosa 563, 600, 601; subdecipiens
567, 600, 606; subdomokosii 582, 592;
626
[Xanthoparmelia] suberadicata 577, 597, 602;
subnigra 563, 566, 600, 601, 606; subochracea
563, 601, 602; subramigera 487; subruginosa
563, 585-586, 601, 602; surrogata 563, 594,
603, 605; synestia 588, 590, 594, 609; tablensis
563, 603, 605; taractica 570, 604; tasmanica
566, 590, 603; terricola 563, 603, 605, 609;
thamnolica 563, 604, 605; toninioides 563, 606,
607; transvaalensis 563, 566, 602, 606, 607;
treurensis 563, 578, 607, 608; tumidosa 563,
602, 607, 608; viridis 563, 590, 609, 610;
wesselsii 563, 609, 610
Xanthoria candelaria 487
Xenasmatella 361, 373; allantospora 361, 365,
372-374; tulasnelloidea 361, 365, 373, 374
Zignoella lichenoides 486; magnoliae 486; nobilis
486
Zoophthora 268; forficulae 263, 268
627
REVIEWERS, VOLUME TWENTY-SEVEN
The Co-Editors express their appreciation to the following individuals who have, prior to
acceptance for publication, reviewed one or more of the papers appearing in this volume.
M. E. BARR BIGELOW R. T. HANLIN G. MORGAN-JONES
T. BARONI D. M. HENDERSON A. ONIONS
J. BOISE L. HOLM S. ONOFRI
P. A. BOWLER J. HOPKINS A. RAMBELLI
M. CHRISTENSEN D. HOSFORD D. J. READ
G. B. CUMMINS R. HUNT R. ROBERTS
R. CURRAH P. W. JAMES J. D. ROGERS
E. E. DAVIS E. B. G. JONES D. J. ROYSE
G. S. DEHOOG B. KENDRICK R. SANTESSON
H. DORFELT R. G. KENNETH N. C. SCHENCK
C. DORWORTH P. KIRK K. SEIFERT
W. M. DOWLER R. E. KOSKE V. SHINDLER
R. S. EGAN P. LANQUETIN R. E. STEVENSON
J. A. ELIX F. F. LOMBARD E. L. STEWART
W. GAMS B. Lowy P. W. TOOLEY
R. L. GILBERTSON N. LUNDQVIST J. M. TRAPPE
M. E. HALE, JR. D. MALLOCH K. WELLS
MYCOTAXON PUBLICATION DATE
Volume 26 (July-September 1986) July 15, 1986
628
ERRATUM, VOLUME TWENTY-TWO
Page 507 line 23 for _ republished
read redrawn to show the hooked paraphyses
ERRATA, VOLUME TWENTY-SIX
Page 33 sn line, 17 for —_7-7-95-A read —_7J-7-85-A
Pages 297-298: These pages were withdrawn by the Managing Editor when
he discovered that the article accepted for those pages by Cryphal, A. P. O.,
entitled “On monotypic and specious genera,” had previously been published
in Phytopathology 46: 411-412. 1956 (cited in Mycotaxon 6: 194. 1977).
ERRATA, VOLUME TWENTY-SEVEN
Page 46 line 33 for Russla read Russula
AS 25 for Stegobolous read Stegobolus
119 22 for Papulasporea parasiticum
read Papulospora parasitica
137 3 for farinacea read farinaceum
263 29 for E. read Entomophthora
35 for E. read Empusa
42 for \.E. read Entomophthora
264 14 for .E read Empusa
266 12 for E. read Empusa
24 for Entomophthora read Empusa
268 iy for £. read Erynia
tse, for E. read Empusa
273 3 for Pseuodparmelia read Pseudoparmelia
287 19 for C. read Claussenomyces
343 9 for ramossissimum read ramosissimum
344 6 for anasaziensis read anasaziense
305 35 for Philippsiella read Phillipsiella
384 48 for fernandezana read fernandeziana
49 for walterii read walteri
409 28 for cinerens read cinereus
436 3 for paracitica read parasitica
459 10 for galapagense read galapagoense
489 25 for guadeloupense read guadelupense
490 25 for granulata read granulatum
507 10 for A, echuyaénsis
read Aecidium echuyaénse
510 1 for echuyaénsis read echyyaénse
2 for echuyaénsis read huyaéns
545 50 for quizotiae read guizotiae
52 for echuyaénsis read echuyaénse
[CONTENTS continued from inside front cover]
ROPES MSEMLINIDIEN Wut ee Aer i) Sac uti Re eee Me Uy Oct Meetn a eee 617
INDe xO -Fungous and Liches axa. >, acs 6% < GS dialeche cain ws vb 4 eh Coe wad 619
EOS ee) AC laerarla nbs WSCA ALN oad GPRM Bb ROMA Oot Sey nC eae Menara eae aie ce 627
Publication date, MYCOTAXON Volume 26... 4.0.06. 2000000200800 0u8 627
a ae EI ee Soames Cpa: Ht at te, 2 RIS she aie Ren.duent a, Maney Os he ee 628
[MYCOTAXON for July-September 1986 (26: 1-520) was issued July 15, 1986]
CO-EDITORS OF MYCOTAXON
G. L. HENNEBERT RICHARD P. KORE
FRENCH LANGUAGE EDITOR ENGLISH LANGUAGE EDITOR
& BOOK REVIEW EDITOR & MANAGING EDITOR
UCL, Place Croix du Sud 3 P.O. Box 264
B-1348 Louvain-la-Neuve, Belgium Ithaca, NY 14851, USA
MYCOTAXON is a quarterly journal devoted to all phases of mycological and lichenological taxonomy and
nomenclature. It seeks to publish all papers within 5 months of submission, using photo-offset lithography.
All articles are reviewed by specialists prior to acceptance. Publication is open to all persons. Papers may be in
French or English, with one or more summaries in any language.
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